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ANNALS OF THE ANNALE VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
VOLUME 94 BAND 94
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
VOLUME 94 BAND
Hee PRUSPEES OF THE DIE SERUSEEBES, VAN DIE
SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM
CAPE TOWN KAAPSTAD
1983-1984
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LISi- OF CONTENTS
Davison, P.
Lobedu material culture: a comparative study of the 1930s and the 1970s. (Published
AUT al SA) ete We ne eames a Sealed Pale a, Baler 10 cytes Jalchig © ag RPRE A MRE eRe ets RAIA oc
ENGLAND, K. W. & Rosson, E. A.
A new sea anemone from South Africa (Anthozoa, Ptychodactiaria). (Published
| SSOP Foes Note Hig re ge hear nen rity ROL UN at a ce MEM Sa Mates a 20 BNE
HeemstraA, P. C. & KANNEMEYER, S. X.
The families Trachipteridae and Radiicephalidae (Pisces, Lampriformes) and a new
Species on A trom! southyAtncas (Rublisheddiune i942) sees eee eee
KANNEMEYER, S. X. see HEEMSTRA, P. C.
Prins, A. J.
A new ant genus from southern Africa (Hymenoptera, Formicidae). (Published
ING@wiembe ral SS) r ets cre Pes cc cccletn sme ie Src PRR teeter nga gh
Prins, A. J.
Morphological and biological notes on some South African arthropods associated
with decaying organic matter. Part 3. The families Dermestidae, Cantharidae,
Melyridae, Tenebrionidae, and Scarabaeidae (Coleoptera). (Published Sep-
LMT Cote SAR) tka toad a a teens eee anne tay gah yack a Ue ARR 2001 heh Mg eo ea
Rosson, E. A. see ENGLAND, K. W.
Volume 94 is complete in 5 parts.
Page
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118)
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NOVEMBER 1983 ISSN 0303-2515
"OF THE SOUTH AFRICAN
/ MUSEUM
CAPE TOWN
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FIsCHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
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Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 94 Band
November 1983 November
Part 1 Deel
Dv HR
A NEW ANT GENUS
FROM SOUTHERN AFRICA
(HYMENOPTERA, FORMICIDAE)
By
A. J. PRINS
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
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OUT OF PRINT/UIT DRUK
ily DO SO), SG, A5 Oo t at), SCS, 5, Oy,
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GDS. ee ICES), LUO), WL SHES), BAG), 23, 35), 250)
EDITOR/REDAKTRISE
Ione Rudner
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ISBN 0 86813 049 4
Printed in South Africa by In Suid-Afrika gedruk deur
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
A NEW ANT GENUS FROM SOUTHERN AFRICA
(HYMENOPTERA, FORMICIDAE)
By
A. J. PRINS
South African Museum, Cape Town
(With 4 figures)
[MS accepted 22 September 1983 |
ABSTRACT
A new genus and its two new species, Agraulomyrmex meridionalis and A. wilsoni, are
described and a key to the workers of the genera of the subfamily Formicinae present in this
subregion is provided.
CONTENTS
PAGE
its SROGHUCTION acs ope eens Rt coe Sy angen on Let ar aaa CYR eet Ete 1
PNEAMLONMDY UITLCX: © CT MO Vine oss oh te, a cncteyeh 9.2, Artois aeegra aun eolepeoee wee 2
VNCAULONMYTINEX METIGIONAUS SP. MOV. ....4-.-4.4+2-4. 08044 seos ne. 2)
PACA ONM TINOGWiSONUSDsMOVas 4.6542 2060s Sees ee 5
Key to the workers of the genera of the subfamily Formicinae in
SOMUMCHMP NU Caden Nam ga wi ate x ena cies ar acd Seo remeunayaace geese 8
PAGKMOMMECSEMENMES) ofr yn kn Gone A Sens 4 ae ayes ne oh nies ean oe 10
NCHS UCINC COMPRIS ACS TCE cate muly dee a lm so elas eee ee Mn Lae sea 11
/: SIDE VIEINOIDS atin os AER aa ee ee een ere OEL ROR Pig Cor Ti ot Iti
INTRODUCTION
In 1969 surveys of ants present in citrus orchards were conducted throughout
South Africa. During the cooler winter months and early spring one sample of
small, brownish hypogaeic ants was collected among some citrus trees in the veld
near Velddrif, a small fishing village about 200 km north-west of Cape Town. As
these ants looked very much like Technomyrmex albipes (F. Smith), which shared
the same habitat, they were noted as such in the collection records. This same
species of ant was also at the time observed in a small citrus orchard near
Citrusdal, about 60 km north-east of Velddrif. In this case they were tending the
aphid Hyadaphis coriandri (Das) on warm days, about 4 to 7 cm above ground
level on the low-growing weeds. They were, however, never observed during the
Summer and autumn months.
Three years later a sample received from the Ministry of Agriculture of
Zimbabwe (formerly Rhodesia) proved to be a different species of this new
genus, which includes the only southern African formicine ants of which the
workers have ten-segmented antennae.
Ann. S. Afr. Mus. 94 (1), 1983: 1-11, 4 figs.
D) ANNALS OF THE SOUTH AFRICAN MUSEUM
Genus Agraulomyrmex gen. nov.
Type species Agraulomyrmex meridionalis sp. nov.
by original designation herein
Diagnosis
Small ants with ten-segmented antennae, flagella incrassate towards apices
without distinct club; first and last segments of flagella longer than wide, the
remaining segments as long as or wider than long. Mandibles triangular with four
to six teeth; frontal carinae short, antennal fossae situated close to posterior
border of clypeus. Eyes large, placed in front of middle of sides of head; ocelli
absent. Maxillary palpi five- to six-segmented, labial palp three- to four-
segmented. Labrum bilobed, widely and shallowly emarginate in middle. Frontal
area clearly indicated.
Alitrunk with promesonotal and mesonotal-propodeal sutures well devel-
- oped, mesometanotal suture absent; propodeum rounded from front to back,
unarmed. Petiole with very small or rudimentary scale overhung by base of
abdomen. Legs moderately long.
Although the ten-segmented antennae point to a myrmelachistine relation-
ship, these ants share characters with the plagiolepidines, viz. the absence of a
distinct antennal club, the structure and form of the alitrunk (the absence of a
distinct metanotum as in certain members such as the genus Acropyga), as well as
the structure of the asepalous proventriculus (Fig. 4A—D). The new genus is
therefore included in the tribe Plagiolepidini.
Workers of these ants bear a close resemblance to those of the genus
Acropyga (Prins 1982); however, they are much smaller and darker in colour, and
the eyes are much larger. In members of Acropyga the eyes are very small,
consisting of only a few facets. The petiolar scale is also much more reduced than
in Acropyga.
Derivation of name
The name Agraulomyrmex (masculine gender) is derived from the Greek
word agraulos, which refers to the mode of living (in the fields).
Agraulomyrmex meridionalis sp. nov.
Description
Worker (Fig. 1A—C)
Specimens collected at Velddrif from which the type material was selected:
TL 2,10-2,20 mm; L0,90-1,0 mm; HL 0,48-0,52 mm; ED 0,26 mm; CL
0,08—0,10 mm; FL 0,40—-0,42 mm; SL 0,32-0,36 mm; WL 0,48-0,50 mm; MFL
0,28-0,30 mm; HFL 0,32-0,36 mm; PL 0,10 mm; CI 76,9-79,2; FI 61,9-65,0; SI
84,2-90,0; CLI 320,0—355,6; TI 58,0-58,3; PI 100,0; HFI 66,7—72,0.
Dark brown, tarsi and mandibles paler. Smooth and shiny all over, especially
declivity of propodeum and petiole. Finely sculptured with piliferous punctures.
A NEW ANT GENUS
Fig. 1. Agraulomyrmex meridionalis sp. nov.
A. Dorsal view of worker. B. Left lateral view of worker. C. Head of worker seen
from the front.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pubescence fairly long, decumbent and yellowish, dense all over except on petiole
and declivity. Pilosity yellowish, long, evident only on mandibles, clypeus,
petiole, and apical margins of abdominal segments.
Head quadrate in dorsal view, about one-fifth to one-sixth longer than wide,
and about one-third wider than pronotum, sides feebly convex, hind margin
almost straight. Clypeus convex in middle, not carinate, its anterior margin
arcuate as in the genus Acropyga. Scapes about two-thirds as long as head
(excluding mandibles) and reaching hind margin; flagellum about two-fifths
longer than scape, first segment as long as the second and third taken together,
apical one slightly longer than three preceding segments taken together; rest as
wide as, or wider than, long. Eyes large, oval, occupying nearly one-fourth of the
length of head (dorsal view—mandibles excluded), and situated in front of middle
of sides. Mandibles shiny, with few large, oval punctures, each bearing erect or
semi-erect seta, and with five to six alternate large and small teeth. Maxillary palp
- six-segmented, labial palp four-segmented.
Alitrunk similar to that of the genus Acropyga, slightly more than three-
eighths longer than wide over pronotum; seen in profile propodeum is on lower
level than promesonotum, declivity almost flat and longer than dorsum of
propodeum. Scale of petiole rudimentary, inclined forward and about as long as
wide and slightly higher than long; seen from behind-almost as wide above as
below, dorsum feebly convex.
Legs moderately long. Abdomen oval in dorsal view. Acidopore on conical
projection of hypopygium, orifice surrounded by fringe of hairs.
Specimens which do not form part of the type material and which were
collected at Citrusdal have the following representative measurements:
TL 2,0-2,20 mm; L 0,94-1,0 mm; HL 0,44—0,48 mm; ED 0,22-0,26 mm; CL
0,09-0,10 mm; FL 0,34-0,39 mm; SL 0,32-0,34 mm; WL 0,46-0,52 mm; MFL
0,26-0,28 mm; HFL 0,32-0,34 mm; PL 0,10-0,11 mm; CI 77,3-83,3; FI 64,1—
68,4; SI 85,0-94,1; CLI 320,0-355,6; TI 53,8-56,5; PI 80,0-90,0; HFI 65,3-69,61.
In most of the specimens examined the head is somewhat shorter than the
alitrunk and the scale of the petiole slightly longer than wide, otherwise similar in
all respects to the Velddrif specimens.
Holotype
1 %, Velddrif, Cape Province (32°47'S 18°10’E), collected by A. J. Prins
7 July 1959, South African Museum specimen.
Paratypes
4 9%, as above, South African Museum specimens.
Other material
Not included in the type material: 4 % 8, Citrusdal, Cape Province (32°37'S
18°58’E), collected by A. J. Prins 17 March 1960, South African Museum
Specimens.
Nn
A NBW ANT GENUS
Derivation of name
The name meridionalis is derived from the Latin word meridional-, which
refers to the southerly distribution of this species.
Agraulomyrmex wilsoni sp. nov.
Description
Worker (Fig. 2A-C)
TL 1.5-1,90 mm; L 0,66-0,7 mm; HL 0,34 mm; ED 0,20-0,21 mm: CL
0,05—0,06 mm; FL 0,28-0,29 mm; SL 0,20-0,21 mm; WL 0,34 mm; MFL 0,14
mm; HFL 0,18 mm; PL 0,07—0,08 mm; CI 88,2—94,1; FI 68,9-75,0; SI 65,6—73,3;
CLI 336,7—440,0; TI 57,8; PI 87,5-114,3; HFI 52,9.
Brown; antennae, mandibles, legs and mesonotum paler in colour, almost
brownish white in some individuals; eyes black. Fairly shiny all over. Finely
sculptured with piliferous punctures as in A. meridionalis, giving integument
an almost fine reticulate or reticulate-punctate appearance; in some specimens
head somewhat duller than in A. meridionalis. Pubescence and pilosity as in
A. meridionalis.
Head quadrate in dorsal view, about one-sixth (or slightly more) longer than
wide and three-eighths wider than pronotum, sides and hind margin almost
straight. Clypeus and mandibles as in A. meridionalis, latter with four sharp
teeth. Scapes slightly more than three-quarters as long as head (dorsal view—
mandibles excluded) and falling short of hind margin by about one-fourth of their
length; flagellum about one-half longer than scape; the first segment as long as
second to fourth taken together; apical one about as long as four preceding ones
taken together; remaining segments as wide as or wider than long. Eyes about
one-fourth the length of the head (dorsal view—excluding mandibles) and
situated in front of middle of sides. Maxillary palp five-segmented, labial palp
three-segmented.
Alitrunk as in A. meridionalis, about three-eighths longer than wide over
pronotum, sutures clearly indicated. Scale of petiole much more reduced, about
as long as wide and about as high as long or slightly higher; seen from behind
almost as in A. meridionalis. Otherwise as in latter species.
Female (Fig. 3A—-B)
Somewhat damaged, both flagella broken off.
TL 3,60 mm; L 1,48 mm; HL 0,48 mm; ED 0,32 mm; LO 0,26 mm; OD 0,13
mm; CL 0,09 mm; FL 0,39 mm; SL 0,34 mm; WL 0,94 mm; MFL 0,26 mm; HFL
Peseern-see 0512 mm; Cl 9157:FL82.1;,S177,3; CLI. 377,8; TI 53,2; Pl 133.3;
HFI 43,6.
Brown, legs, antennae and mandibles paler in colour, eyes black, moderately
shiny. Finely and superficially punctate or reticulate-punctate, especially on legs
and antennae. Pubescence and pilosity as in worker, except for some pilose hairs
on scutellum.
ANNALS OF THE SOUTH AFRICAN MUSEUM
B. Left lateral view of worker. C. Head of worker seen
Fig. 2. Agraulomyrmex wilsoni sp. nov.
A. Dorsal view of worker.
from the front.
A NEW ANT GENUS
mM
Fig. 3. Agraulomyrmex wilsoni sp. nov.
A. Dorsal view of female. B. Left lateral view of female.
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
Head quadrate in dorsal view, nearly one-sixth longer than wide, and slightly
narrower than truncus; sides and hind margin almost straight. Frontal carinae,
fontal area, mandibles, and clypeus as in worker. Eyes large, occupying about
one-fourth the length of the head (dorsal view—mandibles excluded). Scapes
about three-fourths as long as head (dorsal view—mandibles excluded) and
falling short of hind margin by about the length of the second segment of
flagellum.
Alitrunk slightly less than twice as long as wide, parapsidal furrows
indicated, scutellum one-fourth the length of the scutum; seen from above
alitrunk appears oval in outline; in profile propodeum slopes down fairly steeply
towards junction of petiole. Latter with almost obsolete scale which is slightly
wider than long and about one-fourth higher than wide. Abdomen in specimen
examined nearly twice as long as head and alitrunk taken together, oval at base.
Legs moderately long, femora somewhat swollen.
These ants are much smaller than A. meridionalis, much paler in colour, and
more repletes were present.
Holotype
1 2, Gwebi, Zimbabwe (17°40’S 30°40’E), collected by K. J. Wilson
December 1972, South African Museum specimen.
Paratypes
12,7 2%, as above, South African Museum specimens.
Derivation of name
This species has been named after Dr K. J. Wilson of the Ministry of
Agriculture, Harare, Zimbabwe (formerly Salisbury, Rhodesia), who collected
the sample.
KEY TO THE WORKERS OF THE GENERA OF THE SUBFAMILY FORMICINAE
IN SOUTHERN AFRICA
Bolton’s (1973) key should be altered as follows to accommodate the genera
of the southern African subregion (south of the Zambesi River). According to his
key the acidopore is not borne on a conical projection of the hypopygium in the
genus Camponotus, and the orifice is usually not surrounded by a fringe of hairs.
However, in some of our southern African species the acidopore is raised to a
certain extent above the surface of the hypopygium on a somewhat conical area,
and is surrounded by hairs on at least the anterior edge of the orifice. This is
particularly the case in some species of the subgenera, Myrmespera, Mayria (see
Prins 1973), Orthonotomyrmex, Myrmopsamma, Myrmopiromis, and even in
some members of Tanaemyrmex. It is definitely on a conical projection of the
hypopygium in this new genus Agraulomyrmex.
A NEW ANT GENUS 1)
Fig. 4. Lateral view of the proventriculus of the workers of three species of plagiolepidine ants,
compared with that of Agraulomyrmex meridionalis sp. nov.
A. Anoplolepis steingroeveri Forel. B. Acantholepis capensis Mayr.
C. Agraulomyrmex meridionalis sp. nov. D. Acropyga arnoldi Santschi.
=
=)
| s | es fo |e
a |
10.
ANNALS OF THE SOUTH AFRICAN MUSEUM
Antennae9- or lO-sesmenteds 0.5 ieee cian areas qd hc ee ea Cae ee 2
Antennae li- ori2-sesmented tc... aos ser ae | earn eee ne ae 3
Antennae 9-seomented a Arh) vee ate to cana reeenena e Aphomomyrmex
Antennae d 0-sesmented’. <2 ee matic tec seeks ae oe eee eee Agraulomyrmex
Antennac li-seomenteds ova. eat aiapyeny ein 2a cen teal ae SeenON ci ene aes Een 4
Antennae 12-sepsmented, 5... 925. as okt ae sos clos oO ae eae cas ace 7
Maxillary palp 6-segmented, labial palp 4-segmented. Eyes distinct and fairly large;
ocellimay be present 25 2 essa tenis on nce Ue eae Coe ere alg ee rt eee 5
Maxillary palp 4- or 5-segmented, labial palp 3-segmented. Eyes minute ....... Acropyga
Propodeum bidentate or bituberculate; petiole usually bispinose or bidentate above,
occasionally only with upper border strongly emarginate.................. Acantholepis
Propodeum unarmed; petiole neither armed nor deeply emarginate above ........... 6
Metanotum small, much less than half as long as the mesonotum, the mesometanotal
suture feeble or obsolete, at least in the minor workers. Propodeum slightly to strongly
convexcandolteniicher thanthemesometanotimn en ae eee ere Anoplolepis
Metanotum at least half as long as the mesonotum. Metanotum usually fairly
prominent and separated from the mesonotum and propodeum by fairly distinct
sutures. Propodeum about as high as or sometimes higher than metanotum.... Plagiolepis
Petiole reduced to an elongate, low node, allowing the gaster to be reflexed over the
alitrunk. Mandibles elongate triangular, broad, apical tooth long. Maxillary palp
S-sesmentedslabialipalp4-sesmented ane ms sari eine eee Oecophylla
Petiole a node or scale, never as above; mandibles not as above. Maxillary palp 6-
segmented, labial palp 4-segmented, or the palp formula rarely reduced to 3,4 or 3,3,
DUEMEVER SA ek be Sate ey dias Ne MS eracis e AON ne es OND cycle ete aces 8
Antennal insertions very close to, or contiguous with, the posterior clypeal margin.
Acidopore borne on a conical projection of the hypopygium, forming a nozzle, the
onicesurrounded|by. a iringe Of hairs eo aoe a a a ee 9
Antennal insertions some distance (usually greater than basal width of scape) behind
the posterior clypeal margin. Acidopore either not borne on a conical projection, the
orifice not surrounded by a fringe of hairs, or orifice borne on a conical projection or
raised above level of hypopygium on a somewhat conical area, without a fringe of hairs
or with at least some hairs on ventral edge; or the acidopore hidden by a projection of
the pySiauMmies 5 oe eel te le Was Be ceie Sines Rieie: alte 52a Ok pens a 10
Dorsum of alitrunk with very coarse setae arranged in distinct pairs. Eyes at or in
frontokmuidlengthiokthe head) 42a secs sae ae eee Paratrechina
Dorsum of alitrunk with fine setae, not definitely paired. Eyes placed behind midlength
Ol the ead aon. SPs es Bie ees ae ae heer ee ee Prenolepis
Anterodorsal pronotal angles usually projecting as spines or teeth, at least strongly
marginate. Propodeum usually bispinose or bidentate; petiole with sharp angles, spines
Orteethtaboves Monomonpliicr. (Mah sen cia ae eae eee Polyrhachis
Anterodorsal pronotal angles rounded, unarmed. Propodeum unarmed although may
be truncate posteriorly. Petiole a node or scale, never with teeth or spines.
POlyimOrphics.\s Peak tees. BAe sata eRe ae ares Bi eee ee Se cis, <)> cers ee Camponotus
ACKNOWLEDGEMENTS
I should like to thank Dr A. J. Hesse, formerly of the South African
Museum, for his comments as well as Dr V. B. Whitehead and Miss
M. Macpherson, of the same Museum, for their advice. I should also like to thank
Prof. H. J. R. Durr, of the University of Stellenbosch, for identifying the aphid
Hyadaphis coriandri (Das).
a
A NEW ANT GENUS 11
REFERENCES
Botton, B. 1973. The ant genera of west Africa: A synonymic synopsis with keys
(Hymenoptera: Formicidae). Bull. Br. Mus. nat. Hist. 27: 319-368.
Prins, A. J. 1973. African Formicidae (Hymenoptera) in the South African Museum.
Description of four new species and notes on Tetramorium Mayr. Ann. S. Afr. Mus. 62:
1-40.
Prins, A. J. 1982. Review of Anoplolepis with reference to male genitalia, and notes on
Acropyga (Hymenoptera, Formicidae). Ann. S. Afr. Mus. 89: 215-246.
ABBREVIATIONS
CI cephalic index (head width/HL x 100)
CL clypeal length
CLI clypeal index (clypeal width/CL x 100)
ED distance between compound eyes
FI frontal index
FL frontal length
HFI hind femur index (HFL/WL x 100)
HFL hind femur length
HL head length
Ie length of anterior margin of clypeus to base of abdomen
MFL middle femur length
PI __ petiolar index (petiolar width/PL x 100)
PL _petiolar length
SI scape index (SL/head width x 100)
SL scape length
TI thoracic index (thoracic width/WL x 100)
TL total length of body
WL length of alitrunk
Ni - : = 7 7
z —
;
S ;
F \ a ME 2
. ‘ P = -
Se
I,
——s G -
‘f _ ,
ft z
= os x 4 y =
f
i a
— iy
( — 4 ae =
> -
m , a
ti,
a 1
zt i 4
2 ;
iS - c : RES 1 =
- : be : *
2 = : =~
~ - - 3
6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
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Synonymy arrangement according to chronology of bibliographic references, whereby
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In describing new species, One specimen must be designated as the holotype; other speci-
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not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
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Capital initial letters
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e.g. *... the Figure depicting C. namacolus ...’; *. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
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Punctuation should be loose, omitting all not strictly necessary
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‘Revision of the Crustacea. Part VIII. The Amphipoda.’
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Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
A. J. PRINS
A NEW ANT GENUS
FROM SOUTHERN AFRICA
(HYMENOPTERA, FORMICIDAE)
QH
1
eerx “Tt 2 JUNE 1984
| ISSN 0303-2515
NH
jen 2 BRE ARIES sy en
tie
ANNALS
"OF THE. SOUTH AFRICAN -
MUSEUM
CAPE TOWN
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2. LAYOUT should be as follows:
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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Koun, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
“TS Wu 06 ‘ZSEET ISN ‘“UMPW “(OZ8T “MleUog) snivisiu9 NZ *q “O
“TS WU 8rZ “6LOLZ-NVS ‘oftuoanlorg “(OZ8T “Mlouog) smmisi4o NZ “q
“TS WU 96 “O8LLZ-WVS ‘aTtuoanlorg “(L681 ‘AqitsO) wniousdjod puapowusaq “VW
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ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 94 Band
June 1984 Junie
Part 2 Deel
nf
a>
THE FAMILIES TRACHIPTERIDAE
AND RADIICEPHALIDAE
(PISCES, LAMPRIFORMES)
AND A NEW SPECIES OF ZU FROM
SOUTH AFRICA
By
P. C. HEEMSTRA
&
S. X. KANNEMEYER
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
THE FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE
(PISCES, LAMPRIFORMES)
AND A NEW SPECIES OF ZU FROM SOUTH AFRICA
By
P. C. HEEMSTRA
J. L. B. Smith Institute of Ichthyology, Grahamstown
&
S. X. KANNEMEYER
South African Museum, Cape Town
(With 11 figures and 2 tables)
[MS accepted 26 October 1983]
ABSTRACT
The trachipterid fishes of South Africa comprise five species: Trachipterus trachypterus
(Gmelin, 1789), T. jacksonensis (Ramsay, 1881), Desmodema polystictum (Ogilby, 1897), Zu
cristatus (Bonelli, 1820), and a new species Zu elongatus. T. nigrifrons Smith, 1956, is considered
a synonym of 7. jacksonensis, and T. woodi Smith, 1953, appears to be a synonym of
D. polystictum. Zu elongatus sp. nov. is described from four specimens (including two adults)
from off the western Cape Province. It differs from Zu cristatus in body depth, size of eye,
number of lateral-line scales and vertebral counts. Zu elongatus probably also occurs in New
Zealand. The rare adult stage of Zu cristatus is described from a specimen captured off Kenya.
Observations on a juvenile specimen of Zu swimming in the ocean are reported for the first time.
A juvenile specimen of Radiicephalus elongatus Osorio, 1917 (the first South African record for
this species) is also described.
CONTENTS
PAGE
NERO GUCHON Aas Areiae ss yniin: 5A ae Mes einem on a cer eee 13
INES TNO CSW ore coc Se Tape Res tre neecae Mee rier 14
Ramilyanachiptendaes.)o% «14 sb esas selesnas one Saeeee ree 14
Key to genera of family Trachiptendae ...=...:--.5--5--- 15
(GenuSsD eSInOdcCiNGn et ts... a has ne tee eee 15
GENUS HU ACHIPICTUS ie stewie caren ee Be On he ee eee 17
GCM SZ ee recent. < MES FER eS SS 23
amMilyaRaGncepWalida: «jai ss Savatage ae nak aeae aes 35
PNCKHOWIEUS CMICTES ors tel ots cys Beweat Piae Ay ere ee aes 37
FIRE TCREIMCES ert iarae nl ie ere sehen deter eye Sen ce Ree eee Si)
INTRODUCTION
As part of the research being done for a forthcoming revision of J. L. B.
Smith’s (1953) book The sea fishes of southern Africa, the trachipterid fishes of
the J. L. B. Smith Institute were examined. In comparing specimens at the South
iS
Ann. S. Afr. Mus. 94 (2), 1984: 13-39, 11 figs, 2 tables.
14 ANNALS OF THE SOUTH AFRICAN MUSEUM
African Museum with the meagre data from the material in the Smith Institute,
the second author (S. X. K.) discovered a new species of the genus Zu. The first
South African record of the related family Radiicephalidae is based on a juvenile
specimen recently collected in an Engels trawl 70 miles south-west of Cape Point.
METHODS
All measurements are from point to point, and those from the front of the
snout were taken from the premaxillary symphysis with the upper jaw retracted.
Standard length (SL) is measured from the front of the snout to the base of the
upper caudal fin rays. Snout to vent distance (SV) is from the front of the snout to
the anus. Head length is from the front of the snout to the posterior margin of the
gill cavity. Snout length is from the front of the snout to the anterior margin of the
orbit. Lower jaw length is from the anterior tip of that jaw to the posterior end of
the articular bone. Maxilla width is the greatest width of the maxilla. The
anteriormost vertebra with a haemal spine is considered the first caudal vertebra.
Body depth is the greatest depth of the body. Pre-anal vertebrae are those
anterior to a vertical line through the anus. Vertebral counts were made from
radiographs or from cleared and stained material. Lateral-line counts begin with
the scale just above the opercle and stop at the end of the vertebral column.
In the lists of material examined, abbreviations are as follows:
BMNH British Museum (Natural History)
NMNZ National Museum of New Zealand
RUSI J. L. B. Smith Institute of Ichthyology
SAM South African Museum
SIO Scripps Institute of Oceanography
USNM US National Museum of Natural History
Family Trachipteridae
Diagnosis
Body elongate, more or less compressed. Dorsal fin origin above or slightly
behind eye; lateral edges of dorsal fin rays with minute spines or prickles (con-
spicuous in prejuveniles but less obvious in adults). Ventral rays of caudal fin
reduced (rudimentary or absent in some species); pelvic fins rudimentary or
absent in adults; in prejuveniles the pelvic fin rays are elongate and the first ray
bears a row of prickles; anal fin absent. Upper jaw highly protrusile. Posterior
vertebrae of adults elongate, two to four times longer than tenth vertebra; ribs
absent. Ink sac absent; swim-bladder rudimentary. Skin of adults with bony or
cartilaginous tubercles and pierced by numerous pores; lateral line with cartilagi-
nous or bony plates.
Remarks
The monotypic family Radiicephalidae was redefined by Harrisson & Palmer
(1968); the remaining families and genera of the suborder Trachipteroidei have
FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE 15
been reviewed by Walters & Fitch (1960). The trachipterid species of the north-
east Atlantic and Mediterranean were revised by Palmer (1961), and those of the
eastern Pacific were treated by Fitch (1964).
Like the related oar-fish (family Regalecidae), the trachipterids are members
of the off-shore epipelagic community (Parin 1968). They have been observed
swimming in a head-up position by means of undulations of the dorsal fin
(Nishimura & Hirosaka 1964). The peculiar skin tubercles and pore system are
thought to reduce drag when the fish is swimming (Walters 1963). Three genera
are recognized. Although more than thirty species have been described, there are
probably fewer than ten valid species in this family.
KEY TO GENERA OF FAMILY TRACHIPTERIDAE
la Caudal fin comprises 4-10 dorsal rays approximately parallel to the long axis of the body;
ventral caudal rays absent; no elongate spiny plates or bony tubercles along ventral edge of
cbt PIE 8 PSS k tks GN. aay 8 es Mee lieg Ia Ps oi hie oop eels cuaiae Agia Desmodema
1b Caudal fin of two parts: 6—12 dorsal rays usually set at a steep angle to the long axis of the
body and 1-5 ventral rays (rudimentary spiny nubbins in Trachipterus and adult specimens of
Zu); elongate spiny plates or bony tubercles along ventral edge of tail.................. 4
2a Lateral line runs along ventral edge of tail as a zigzag series of elongate spiny plates; subadults
(less than 700 mm SL) with ventral edge of body distinctly scalloped; body depth 1,5—2,9 in
PON MIE ie eR et dealin ns AAG wale Sidwed Vek oe hes PSSA ME ame yeu ee. eee Aen Zu
2b Lateral line straight, well above ventral edge of tail in adults (specimens more than 1 500 mm
SL); ventral edge of tail in adults with bony tubercles but no elongate spiny plates; ventral
cancomnoay straight: body depth 3;7—4,lin SV ..-.. 2.52 52220: +s8- ens ee: Trachipterus
Genus Desmodema Walters & Fitch, 1960
Diagnosis
Seven dorsal pterygiophores before first neural spine; no predorsal bones;
caudal fin on same horizontal axis as caudal peduncle, comprising 4—10 dorsal
rays; ventral caudal rays absent; tail greatly elongated in juveniles and adults;
ventral body profile of prejuveniles (specimens less than 100 mm SV) not scal-
loped. Prejuveniles silvery, with numerous round dark spots.
Remarks
This genus was recently revised by Rosenblatt & Butler (1977). They recog-
nize two species. D. Jorum Rosenblatt & Butler, 1977, is known only from the
North Pacific.
Desmodema polystictum (Ogilby, 1897)
Fig. 1A
Trachipterus jacksoniensis polystictus Ogilby, 1897: 649.
Trachipterus sp.: Smith, 1949: 504, fig. 264b.
Trachipterus woodi Smith, 1953: 504, fig. 264b; 1966: 83, pl. 15 (fig. A).
Trachypterus woodi: King & Ikehara, 1956: 22, fig. 5.
Desmodema polystictum: Rosenblatt & Butler, 1977: 848, figs 1-4.
(See Rosenblatt & Butler (1977) for additional synonyms and references.)
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
Material
Holotype of 7. woodi Smith, RUSI 133, SV 89 mm (rear end of tail missing);
Xora River mouth, Transkei.
SIO 75-139, SL 100 mm, SV 72 mm; 10°00’N 119°00’W; midwater trawl;
0-50 m.
SAM-—27786, SL 96 mm, SV 68 mm; found in a tidal pool at Simon’s Town,
False Bay, Cape Peninsula.
USNM 164171, SL 113 mm, SV 78 mm; 04°41’'N 159°53’W (radiograph only
seen).
Diagnosis
This diagnosis is according to Rosenblatt & Butler 1977, plus data from
specimens mentioned above.
Dorsal fin rays 120—128; pectoral fin rays 12-14; caudal fin rays 7-10; gill-
rakers 2-3+9-10. Vertebrae: 71-74 total, 18-20 precaudal, 37—42 pre-anal.
Eye diameter greater than snout length. Tail greatly elongated in adults; SV
about two-thirds SL in prejuveniles and one-third SL in adults.
Remarks
Smith (1966) gave an amended, amplified description of the holotype of
T. woodi. The following data not supplied by Smith are added here. SV 89 mm,
body depth at pelvic fin origin 33 mm, head length 24mm, eye diameter
10,1 mm, snout length 7,0 mm, maxilla width 6,0mm, lower jaw length
10,6 mm. All the pelvic fin rays have been broken off at the base, but there seem
to be 7 nubbins on the left side and 8 on the right. Because of the poorly ossified
bones of this small specimen (perhaps made worse by preservation in formalin), it
is difficult to get a clear radiograph. The anterior dorsal fin pterygiophores in
relation to the first neural spine and the transition from precaudal to caudal
vertebrae cannot be discerned, but the number of pre-anal vertebrae is clearly 41.
Data from the SAM specimen are as follows: SL 96 mm; SV 68 mm; dorsal
fin rays 128, first five rays elongate, the longest is 37 % SL; pectoral rays 13; pelvic
rays 9, elongate, reaching caudal fin; caudal rays 8; gill-rakers 3+ 9; lateral-line
scales 158, each plate with a single spine except fifteenth, which has a double
spine; anus asymmetrical, on left side 1,5 mm above ventral edge of body.
Vertebrae: precaudal 20, pre-anal 37, total 74.
Comparison of the holotype of T. woodi with one of the prejuvenile speci-
mens of D. polystictum examined by Rosenblatt & Butler (1977) revealed no
significant differences between these two specimens. 7. woodi is therefore con-
sidered a synonym of D. polystictum. The possibility of T. woodi being a synonym
of D. polystictum and the record of T. woodi from the Pacific (King & Ikehara
1956) were apparently overlooked by Rosenblatt & Butler (1977).
Distribution
South Africa, Australia, Japan, eastern Pacific and western Atlantic.
FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE 17
Genus Trachipterus Gotan, 1770
Diagnosis
Body depth of adults 3,7—4,4 in SV, not greatly constricted behind vent.
Ventral edge of body straight, with enlarged pointed tubercles. Lateral-line plates
of adults more or less fused to form an almost continuous tube with rudimentary
blunt spines; prejuveniles and juveniles with separate, oval lateral-line plates,
each with a central, well-defined spine. In adults the lateral line runs well above
the ventral edge of the tail, but in smaller specimens the lateral line runs fairly
close to the ventral edge of the tail along the rear half of the tail. Dorsal and
pelvic fin rays of prejuvenile stage without series of bulbous or leaf-like appen-
dages; first five or six dorsal rays, pelvic and caudal rays elongate in prejuveniles
and juveniles. Dorsal caudal rays well developed, set at steep angle to horizontal
axis of peduncle. One pterygiophore before first neural spine, and 8 or 9 between
first and second neural spines. Scales absent.
KEY TO SOUTH AFRICAN SPECIES OF TRACHIPTERUS
la Pectoral fin rays 8-11; eye diameter 1,4-1,6 in lower jaw length; tubercles small and
indistinct except along ventral edge of body and tail, not arranged in rows along each side of
SsiiSRPECRYUSIOPHORES 5 2. Gs. eeu gids Sean ake ote ere eee meee Welee T. trachypterus
15 Pectoral fin rays 13 or 14; eye 1,9-—2,2 in lower jaw length; tubercles distinct, those along
ventral edge of body and tail larger, pointed and slightly curved; tubercles arranged in rows
alonmeaen side of dorsal fin pterypiophores ..........2..26.2..5-2---=- T. jacksonensis
Trachipterus jacksonensis (Ramsay, 1881)
Figs 2-3
Regalaecus jacksonensis Ramsay, 1881: 631, pl. 20.
Trachipterus jacksonensis: McCulloch, in Whitley, 1927: 296, pl. 25 (fig. 2).
Trachipterus nigrifrons Smith, 1956: 449, text fig. and pl. 12; 1965: 18.
Material
Holotype of Regalaecus jacksonensis Ramsay, Australian Museum A9114
(radiograph only).
Holotype of T. nigrifrons Smith, RUSI 52, 1 210 mm total length (fide Smith
1956), East London.
RUSI 13128, 1 320+ mm total length; East London.
SAM-—25093, 2 110 mm SL, 1 050 mm SV; trawled west of Cape Town.
SAM-—26731, 1 220+ mm SL, 600 mm SV; trawled west of Cape Peninsula.
SAM-—28380, 1 490 mm SL, 700 mm SV; trawled 34°31’S 17°59’E.
Diagnosis
Dorsal fin rays 166-173, the first 6 rays short and hairlike; pectoral fin rays 13
or 14; gill-rakers 3-5+8. Vertebrae: 81-83 total, 31—34 precaudal, 44—48 pre-
anal. Eye diameter 1,9—2,2 in lower jaw length, 6,0—7,3 % SV. Dermal tubercles
on body distinct, those along ventral margin of body and tail enlarged, pointed
and slightly curved; tubercles in rows along each side of dorsal fin pterygiophores
18
ANNALS OF THE SOUTH AFRICAN MUSEUM
Drawn from holotype of T. nigrifrons Smith, RUSI 52, 1 210 mm TL. Pectoral
f anus; inset shows detail of skin tubercles from middle of tail
fin reconstructed. Arrow indicates position o
Fig. 2. Trachipterus jacksonensis (Ramsay, 1881).
FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE 19
Fig. 3. Trachipterus jacksonensis (Ramsay, 1881). Arrangement of tubercles in
vertical rows along dorsal fin pterygiophores.
(Fig. 3). Head and body silvery; front of head, interorbital area, dorsal part of
upper jaw and tip of lower jaw black.
Remarks
Smith (1956) compared his new species 7. nigrifrons with two species
(T. ishikawae and T. ijimae) described by Jordan & Snyder (1901) from Japan.
T. ijimae was synonymized with Zu cristatus by Fitch (1964: 231). In comparing
T. nigrifrons and T. ishikawae, Smith (1956) used data from a later paper by
Jordan & Snyder (1904), which is based on two non-type specimens that may or
may not be conspecific with the holotype of T. ishikawae. In the original descrip-
tion of T. ishikawae the numbers of pectoral fin rays and gill-rakers are not given,
and without having recourse to the holotype the identity of this species must
remain in coubt.
Smith (1956) apparently overlooked the detailed description of T. jack-
sonensis (Ramsay, 1881) given by McCulloch (in Whitley 1927). McCulloch’s
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
description agrees with Smith’s (1956) account of T. nigrifrons, and McCulloch,
after comparing his specimen with the holotype of T. jacksonensis, stated that
(p. 297) ‘it appears a little deeper than the holotype, to which it is exactly similar
in every other detail’.
Distribution
South Africa and Australia.
Trachipterus trachypterus (Gmelin, 1789)
Fig. 4
Cepola trachyptera Gmelin, 1789: 1187.
Trachypterus arawatae Clarke, 1881: 195, text-fig.
Trachipterus jacksonensis (non Ramsay): Hamilton, 1916: 372, fig. 1.
Trachipterus arcticus (non Briinnich): Barnard 1925: 353,; pl. 14 (fig. 3); 1948: 359, fig. 17.
Trachipterus iris: Smith, 1949: 141, fig. 264.
Trachipterus trachypterus: Hamilton, 1916: 374, figs 3-6. Palmer, 1961: 342, pl. 62 (fig. 2).
(See Palmer (1973) for additional synonyms and references.)
Material
BMNH 1891.8.31.27-35, 7 specimens, North Atlantic (radiograph only
seen).
RUSI 13106, 1 620+ mm SL, 710 mm SV; South Africa, no data.
SAM-=11894, 540+ mm SL, 236 mm SV; South Africa, no data.
SAM-22078, 1 860 mm SL, 900 mm SV; trawled off Dassen Island (33°25'S
18°E).
SAM-—23677, 113 mm SL, 82 mm SV; trawled off Slangkop, Cape Peninsula.
SAM-—24024, c. 520 mm SL (damaged), 250 mm SV; trawled west of Cape
Town.
SAM-—26708, 55 mm SL, 40 mm SV; 33°06’S 16°08'E; 150-0 m; N100B
plankton net.
Diagnosis
Dorsal fin rays 145-184, first 5 or 6 rays elongate in juveniles and prejuve-
niles, forming a dorsal pennant; pectoral rays 9-11; gill-rakers 3-5 + 10. Verte-
brae: 84-96 total, 35-39 precaudal, 49-57 pre-anal. (Vertebral counts for the
New Zealand specimen NMNZ 2056 are c. 97 total and about 42 precaudal.) Eye
diameter (of adults) 1,4—1,6 in lower jaw length, 8,4-8,9 % SV. Dermal tubercles
on body small, except along ventral margin of body and tail, uniformly distributed
over dorsal fin pterygiophores (Fig. 4). Head and body silvery; juveniles with 3-5
dark blotches on body.
Remarks
The present specimens generally agree with Palmer’s (1961) account of
T. trachypterus (Gmelin, 1789), except that they do not have the body axis curved
dorsally in the posterior caudal region, and the 540mm SL specimen
FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE 21
Fig. 4. Trachipterus trachypterus (Gmelin, 1789). Uniform arrangement of tuber-
cles along dorsal fin pterygiophores.
(SAM-11894) seems to have the greatest body depth nearer the anus than the
head, instead of just behind the head. According to Palmer (1961) T. trachypterus
differs from 7. arcticus in the dorsally curved tail and in having the greatest body
depth immediately behind the head; in T. arcticus, the tail is straight and the
greatest body depth is ‘3 to 3 of the way along its length, except in specimens of
less than 300 mm’.
The original description of Cepola trachyptera Gmelin, 1789, is inadequate to
distinguish which species Gmelin was trying to describe: ‘C. capite declivi, maxilla
utraque fornicata, pinnis aculeatis serratis scabris. Habitat in mari adriatico, linea
laterali recta, squamarum serie simplici; an secundum Forsterum proprium genus?’
(Cepola with downward-sloping head [i.e. steep forehead], maxilla arched on either
side, fins rough [with] saw-like prickles. Habitat in Adriatic Sea, lateral line straight,
scales in a single row; according to Forster it may belong in a genus of its own.)
C. trachyptera was the third species of Cepola listed by Gmelin (1789: 1186-
1187); the first two (C. taenia and C. rubescens) are considered synonyms of
pip ANNALS OF THE SOUTH AFRICAN MUSEUM
C. macrophthalma (Linnaeus, 1758). By his placement of trachyptera in Lin-
naeus’s genus Cepola, Gmelin probably recognized at least a superficial similarity
between his new species and C. macrophthalma. Hence it seems reasonable to
suppose that C. trachyptera was an elongate fish with a markedly compressed
body and a dorsal fin that ran the length of the body. The steep forehead rules out
trichiurids.
Except for trachipterids, the only Adriatic fishes with an elongate com-
pressed body, steep forehead, and long dorsal fin are the oarfish Regalecus glesne
Ascanius, 1772, and Lophotus lacepedei Giorna, 1809. The statement about the
fins ‘pinnis aculeatis serratis scabris’ apparently refers to the row of minute spines
or prickles along the lateral edges of the dorsal fin rays of trachipterid fishes. The
species name trachyptera means ‘rough fin’, being derived from the Greek words
trachys (=rough) and pteron (= wing or fin). These fin-ray prickles are lacking in
the oarfishes but very conspicuous in juvenile trachipterids.
We are left, then, with Trachipterus trachypterus (sensu Palmer 1961) and Zu
cristatus as the Adriatic species that best fit the description given by Gmelin.
Because of its greater body depth (which would render the ‘Cepola’ designation
less likely) and the distinctive features of the prejuveniles (which are unlikely not
to be noticed and mentioned in a description), Zu cristatus seems the less
plausible of the two species. Furthermore, 7. trachypterus is fairly common in the
Adriatic Sea (Jardas 1980).
By a process of elimination together with a few assumptions, we have come
to the conclusion that Cepola trachyptera Gmelin is the species currently recog-
nized as Trachipterus trachypterus.
Hamilton (1916) published descriptions of seven specimens of trachipterids
from New Zealand. The specimen from Nelson, South Island, that he illustrated
(fig. 2) from his original sketch was doubtfully assigned to T. jacksonensis. This
specimen, later illustrated by Whitley (1968: 46, fig. 1) under the name ‘Des-
modema arawatae’, appears to be Zu elongatus sp. nov. Judging from the count
of eleven pectoral fin rays given for five of the specimens, Hamilton’s other
specimens appear to be TJ. trachypterus. Hamilton identified his largest,
‘approximately 6 ft’ specimen with T. jacksonensis because it had smooth dorsal
fin rays, instead of ‘all rays spinous, and having a spiny tubercle at the base’ for
his juvenile specimens of T. trachypterus. Hamilton assumed that the minute
spinules or granulations on the dorsal fin rays of juveniles were a good specific
character, and were not lost or greatly reduced with growth. However, this is
precisely what happens with all trachipterids. In our adult specimens of
Trachipterus and Zu the spinules are absent or very much reduced, compared
with those of juveniles.
Distribution
Mediterranean Sea, eastern Atlantic to South Africa, Japan and New Zea-
land.
FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE 23
Genus Zu Walters & Fitch, 1960
Diagnosis
Young with pelvic and anterior 6 dorsal fin rays greatly elongated, bearing
serially arranged, membranous flaps (Figs 1D, 8). One pterygiophore before first
neural spine, and 8 or 9 between first and second neural spines. Scales present,
though very deciduous, and not apparent in prejuveniles. Body depth 1,7—2,8 in
SV, more or less constricted behind anus. Ventral edge of body distinctly scal-
loped in specimens less than 700 mm SL; belly of specimens more than 800 mm
SL nearly straight, but with a low fleshy keel from pelvic fin rudiments to anus.
Lateral line drops to ventral edge of tail shortly behind anus, from where it runs
aS a zigzag series of elongate spiny plates to base of caudal fin and is then
continued on to the ventral caudal filament (of prejuveniles) as a straight tube.
Total vertebrae 63-69 or 84-87.
KEYolO;SPECIESIOE ZU
la Body depth 12-16% SL; prejuveniles with body not abruptly constricted behind vent; eye
phere lateral-linesplates 126-130 2... ce. ne a le ee Z. elongatus sp. nov.
1b Body depth 20-26% SL; prejuveniles with body abruptly constricted behind vent; eye
easier orsy « lateral-line plates 99-106... a... ccs ae sees aa eee Z. cristatus
Zu cristatus (Bonelli, 1820)
Figs 1B—D, 5-8
Trachypterus cristatus Bonelli, 1820: 487. Smith, 1949a: 99; 1949b: 142, fig. 264a. Dieuzeide et
al., 1954: 151, figs. Tortonese, 1958: 1. Parin et al., 1978: 176.
Gymnetrus repandus Metaxa, 1833: 53, fig. 7.
Trachypterus bonelli Valenciennes, in Cuvier & Valenciennes, 1835: 331.
Gymnetrus miillerianus Risso, 1840: 13, pl. 125.
Trachypterus repandus: Costa, 1842: 11, pl.
Trachypterus semiophorus Bleeker, 1868: 279, pl. XII. Weber & de Beaufort, 1929: 89.
Trachypterus iris (non Walbaum): de Buen, 1917: 23, fig.
Trachypterus gavardi Bounhiol, in Bounhiol & Gavard, 1923: 1.
Trachypterus ijimae Jordan & Snyder, 1901: 311, pl. 17 (fig. 9). Tanaka, 1915: 358, pl. 96 (fig.
301). Bolin, 1933: 35. Okada & Suzuki, 1956: 168, fig. 1. Lindberg & Legeza, 1965: 298.
Zu cristatus: Walters & Fitch, 1960: 445. Palmer, 1961: 346. Fitch, 1964: 229, fig. 1. Backus er al.,
1965: 139, fig. 2. Ibanez & Gallego, 1974: 523, fig. 1. Karrer, 1975: 72. Fitch & Schultz, 1978:
89, fig. 5. Parin et al., 1978: 176. Oliver, 1982: 60, figs 1-2.
(See Palmer (1973) for additional references.)
Material
RUSI 7185, 2 specimens: 64 mm SL, 31 mm SV; 204 mm SL, 90 mm SV;
South Africa, no data.
RUSI 7187, 232 mm SL, 106 mm SV; Algoa Bay, eastern Cape Province.
RUSI 7188, 173 mm SL, 77 mm SV; Kariega River mouth, 33°36'S 26°42’E.
RUSI 13352, 950 mm SL, 450 mm SV; off Mombasa, Kenya.
SAM-—24522, 415 mm SL, 170 mm SV; off Slangkop lighthouse, Cape Penin-
sula, Cape Province.
SAM-—27679, 248 mm SL, 107 mm SV; 28°23,5’S 32°41,9’E.
ANNALS OF THE SOUTH AFRICAN MUSEUM
24
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‘¢ “314
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FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE
‘sajejd oul] [e19je] JO saseq BULIDAOD AqjeIIed sayeog “6L9LZ-WVS ‘oytuoanloig
‘(OZ8I
6
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ANNALS OF THE SOUTH AFRICAN MUSEUM
26
‘snue
puryaq ysnf poyenqis ‘poyoryje juowrsid YIM sojeos uTYy) AIOA Jo sayojed Aulys “6L9LZ-WVS ‘oTtueanfoig “(OZ8T ‘I[ouog) sniwisiua NZ */ ‘BLA
FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE a
Diagnosis
D 120-150; P 10-12; V 5-7 (absent in adults); dorsal caudal fin rays 6-12,
set at a steep angle to horizontal axis of caudal peduncle; ventral caudal rays
represented by spiny nubbins in adult; prejuveniles with 1—5 ventral caudal rays,
with 1 or 2 produced into long filaments. Gill-rakers 2-3+8-9. Lateral-line
plates 99-106; lateral line begins above middle of eye and continues beyond the
vertebral column on to the caudal fin between the second and third ventral caudal
rays (about 16 pored scales on the caudal membrane of SAM-—27679). Vertebrae:
63— 69 total, 22—24 precaudal, 32 pre-anal. SV 41-48 % SL; body depth 20-26 %
SL; eye diameter 1,3—1,7 in length of lower jaw, 13-16% SV. Young silvery,
with about 6 wavy dark vertical bars on dorsal part of body and 4 on ventral part;
tail with about 6 complete black bars; caudal fin mostly blackish, proximal part
pale. Adults silvery grey, paler ventrally; dorsal fin scarlet; caudal fin reddish
black, darker distally.
Description of adult (RUSI 13352, 950 mm SL)
D 145; P 11; dorsal caudal fin rays 9; gill-rakers 3+ 8. Body robust (not
nearly as compressed as adults of Trachipterus or Zu elongatus sp. nov.), with
fleshy keel along belly; body covered with small, very deciduous, thin, cycloid
scales. (All scales on body are missing, but the scale pockets are distinct; a few
scales were left on the tail near the lateral line.) Lateral line curving gradually
from above opercle to ventral edge of tail 100 mm behind anus; lateral-line scales
spiny, represented by two rows of staggered spiny scutes along ventral part of tail.
Pelvic fins represented by a few nubbins. Single nostril on each side of snout.
Pseudo-branch well developed. Pyloric caeca very numerous. Swim-bladder not
apparent.
Measurements for six specimens are given in Table 1.
TABLE 1
Measurements (in mm) of specimens of Zu cristatus.
RUSI RUSI RUSI RUSI SAM SAM
7185 7187 7188 13352 24522 27679
Standard length (SL) 205 232 173 950 415 248
Snout to vent (SV) 91 108 76 451 170 107
Greatest body depth 49 56 43 223 84 64
Body depth at anus 19 21 18 123 4] 23
Head length 31 — 7a 161 60 SH
Eye diameter 12 14 10 69 Di 14
Maxilla width 6,3 4.3 5,8 By 18) 9
Lower jaw length 19 22 16 92 36 8,
Remarks
Adults of Zu cristatus are apparently rarely captured; only four specimens
longer than 800 mm SL have been reported (Bolin 1933; Tortonese 1958; Ibanez
& Gallego 1974; Fitch & Schultz 1978). Judging from the illustration of a 610 mm
ANNALS OF THE SOUTH AFRICAN MUSEUM
28
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ajiuaanf aaty e Jo SuIMLIG “YISUIT UI WU QQT NoGY “(OZ8T ‘]aUOg) snwISU9 NZ °g “314
aca
| | i 4
— — ' «1200IEN S EE OO OOOO.—OOOOanawnaawn nn I.
FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE 29
prejuvenile by Tortonese (1970, fig. 190b), metamorphosis takes place in the
600-800 mm size range. The small prejuveniles that we have examined
(64-248 mm SL) are much more compressed than the adults. The pelvic fins and
elongated anterior dorsal fin rays are lost during the transition from prejuvenile to
juvenile stage.
Despite previous reports of deciduous cycloid scales on specimens of Zu
cristatus (Bolin 1933; Tortonese 1958; Walters & Fitch 1960; Palmer 1961; Fitch
1964), Rosenblatt & Butler (1977) discounted these observations, because they
could not find scales on their eight specimens (27,5—-811 mm SL). As noted
above, adults of Zu do have scales but the epidermis is very fragile, and there are
only a few scales remaining on the tail of the 950 mm SL specimen. On the
248 mm SL prejuvenile (SAM-—27679), the scales are most apparent along the tail
(Fig. 5), which they appear to cover completely, and continue on to the caudal fin
rays. Some of the bases of the lateral-line plates are partially covered by scales
(Fig. 6). The lateral surface of the tail just behind the anus has scales that seem to
have pigment attached to them. These scales are apparent on the fish as large
shiny patches (Fig. 7). The scales forming these shiny patches are exceedingly
thin, but will separate on being lifted with a fine needle. There is evidence of
Squamation over the entire body area, particularly ventrolaterally where many
scaled areas remain intact. The head does not appear to have scales.
Except for the report of Nishimura & Hirosaka (1964) of a large Trachipterus
specimen in a relatively small aquarium, there have been no published observa-
tions on live trachipterids. In view of the dearth of such reports, the following
observations of L. Jones of Fynnlands, Natal are of considerable interest. Judging
from Jones’s sketch, the specimen that he observed was probably Zu cristatus.
‘I was swimming just beyond the surf-line in 20 feet of water when I noticed
what at first appeared to be a small fish caught by a stinging plankton or jellyfish
some 6-10 inches below the surface. On closer examination I found it to be a fish
with long thread-like fins: . . . a small silvery fish with big head and eye, with red-
brown markings, about 33 inches long. The six [elongated] anterior dorsal rays
were at least 15 inches long, with what appeared to be pairs of tiny brown leaf-like
appendages set 13 to 2 inches apart down the length of each dorsal ray. Its pelvics
were also very long, about the same length as the body, but I can’t remember if
the “‘leaves”’ were present on these or not. Its tail was all but invisible, but the
dorsal fin seemed to be part of its propulsive mechanism (the lower or rear part
that is) for it swam in a head-up-tail-down manner which I have drawn for you’
(Fig. 8).
Zu elongatus sp. nov.
Figs 9A-11B
? Trachipterus jacksonensis (non Ramsay): Hamilton, 1916: 374, fig. 2.
Trachipterus jacksoniensis (non Ramsay): Phillipps, 1944: 120, pl. 52. (Unjustified emendation. )
Trachypterus arcticus (non Briinnich): McCann, 1953: 5 (figs 5—6 only).
? Desmodema arawatae (non Clarke): Whitley, 1968: 45, fig. 1.
ANNALS OF THE SOUTH AFRICAN MUSEUM
30
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WIOl
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FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE 31
Holotype
SAM-—24704, 1 166 mm SL, 404 mm SV; adult, ?¢. Trawled south-west by
west of Cape Columbine, north-western Cape Province; bottom depth 411 m;
donated by Captain Jacoby, S. T. Rochea; collected 22 November 1965.
Paratypes
SAM-—29392, 1 135 mm SL, 421 mm SV; adult, ?¢. Trawled west of Cape
Town by Irvin & Johnson Ltd trawler; depth and date of collection unknown.
SAM-—19870, 630+ mm SL (part of tail missing), 228 mm SV; juvenile.
Trawled north-west of Table Bay by trawler of Irvin & Johnson Ltd; bottom
depth 580 m; collected in 1946, exact date unknown.
SAM-—24414, 480 mm SL (upper jaw fully protracted), 166 mm SV; prejuve-
nile. Trawled off Cape Columbine, north-western Cape Province; bottom depth
548 m. Collected and donated by P. Henry, Midwest Fisheries Ltd, Cape Town, 1
February 1965.
Other material
SAM—29396, 1 123 mm SL, 421 mm SV; adult, badly damaged; no data.
NMNZ P.834, c. 425 mm SL, 138 mm SV; French Pass, New Zealand.
NMNZ P.2248, 315 mm SL, 108 mm SV; Awanui, New Zealand.
Diagnosis
D 142-147; P 11-12; V 7, elongate in young, absent in adults; C 17 (12
dorsal rays and 5 ventral rays of which 2 may be represented by tiny spines). Gill-
rakers 2—3+ 7-9. Lateral-line plates 127-130. Vertebrae: 84—87 total, 29-31
precaudal, 38—40 pre-anal. SV 34-37 % SL; body depth 12-16 % SL; eye 9-10 %
SV. Teeth: 9-21 in upper jaw, 6—9 in lower law; adults with 2 vomerine teeth;
palatine teeth present or absent. Dermal tubercles and pore system present at all
stages; scale pockets well developed in adults (Fig. 10).
Description (measurements in Table 2)
Adult (holotype, 1 166 mm SL; Fig. 9A)
Body laterally compressed, tapering gradually to the elongate tail; dorsal
body profile convex, the ventral profile irregular; fleshy keel along ventral midline
of belly. Body, tail, and cheeks covered with tubercles and conspicuous pores;
scale pockets distinct; tubercles on cheek enlarged.
D 143, the posterior rays long, filamentous; P 11; pelvic fins absent, repre-
sented by bony ridge within a slit-like pocket; 2 of the 5 ventral caudal fin rays are
small rudimentary spines; the other caudal rays elongate, the longest 160 mm.
Gill-rakers 3 + 8; lateral-line plates c. 128. Teeth: upper jaw with 10 on right side
of symphysis, 11 on left side; lower jaw with 6 teeth on right side of symphysis and
3 on left side; 2 teeth on vomer, 3 teeth on right palatine, 2 teeth on left palatine.
Colour in alcohol: body tan, interspersed with paler tubercles; front of head
with triangular dark area, extending on to lips and gular region; posterior dorsal
fin rays black; caudal fin black except for a small, pale area proximally.
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Zu elongatus sp. nov. Paratype, adult, SAM—29392, 1 135 mm SL. Scale
pockets.
TABLE 2
Measurements (in mm) of specimens of Zu elongatus sp. nov. Specimen SAM-24414 was fixed
with the upper jaw completely protracted and the head is somewhat damaged, consequently the
measurements SL, SV and head length are approximate estimates of what they would be with the
upper jaw retracted. Both eyes are also missing on this specimen.
Holotype Paratypes
SAM Catalogue No. 24704 19870 24414 29392 29396
Standard length (SL) 1166 627+ 460 1168s 1123
Snout to vent (SV) 404 228 154 421 421
Greatest body depth Wa 99 63 177 176
Body depth at anus 1138) 66 34 121 13%
Head length 137, 81 66 134 135
Eye diameter 4] 7h — 42 42
Maxilla width 38) 16 13 33 36
Lower jaw length 73 42 34 74 80
FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE 33)
Prejuvenile (480 mm SL; Fig. 11A)
Body elongate, laterally compressed, gradually tapering to caudal fin, not
abruptly constricted behind vent; dorsal profile of body convex; ventral profile
anterior to anus strongly scalloped, with three well-defined ventral lobes. The
strong upward curve of the tail, characteristic of prejuveniles in Zu cristatus, is
not apparent in this specimen. Lateral line curving gradually from above opercle
to ventral margin of tail 30 mm behind anus. Body surface abraded; no scales or
scale pockets apparent.
D 147, the first 6 rays elongate; P 11; pelvic fins elongate; caudal directed
upward, the rays elongate (longest 170 mm). Gill-rakers 3+ 8-9; lateral-line
plates c. 127. Teeth: upper jaw with 8 on right side of symphysis, 10 on left side; 4
on each side of lower jaw symphysis; no teeth on palatines or vomer.
Colour in alcohol: three incomplete dark cross-bars on body; five dark bars
on tail. Triangular dark area above and extending across symphysis of upper jaw;
dusky area across lower jaw symphysis, extending on to gular region. Distal part
of caudal fin dark; some dorsal fin rays with traces of dark pigment.
Juvenile (630+ mm SL; Fig. 11B)
Body shape as for prejuvenile; ventral lobes smaller and less distinct. Caudal
fin missing; anterior dorsal rays and pelvic fins elongate. Dermal tubercles on
cheek enlarged; scale pockets visible at 40 x magnification.
D 137; P 12; gill-rakers 2-3 +7; lateral-line plates c. 130. Teeth: upper jaw
with 7 on right side of symphysis, 4 on left side; lower jaw with 2 teeth on right
side and 4 on left; 2 teeth on vomer and 1 on right palatine.
Colour in alcohol: six dark cross-bars on tail; symphysis of lower jaw dark.
Body abraded, with no other pigmented areas apparent.
Paratype SAM-—29392, 1 135 mm SL (Fig. 9B), differs from the holotype
SAM-—24704, 1 166 mm SL and from the other adult, SAM—29396, 1 123 mm SL,
in that it lacks palatine teeth. SAM-—29396 has only 9 teeth in the upper jaw.
However, the teeth in Zu elongatus are easily detached and since SAM-29396 is a
badly damaged specimen some teeth might have been lost.
Remarks
As denoted by the name, Zu elongatus is more elongate than Zu cristatus
(body depth 12-16% SL, versus 20-26 % SL). Other differences between these
two specimens are the size of the eye (9-10 % SV in Zu elongatus, 13-16 % SV in
Zu cristatus), number of the lateral-line plates (126-130 in Zu elongatus, 99-106
in South African specimens of Zu cristatus, but 114-118 reported by Fitch (1964)
for eastern Pacific material), and vertebral counts (84-87 total, 29-31 precaudal,
38—40 pre-anal for Zu elongatus; 63-69 total, 22-24 precaudal, 32 pre-anal for Zu
cristatus). Palmer (1961) and Fitch (1964) give counts of 64-65 and 63 for total
vertebrae of Zu cristatus from the eastern Atlantic and eastern Pacific. The ranges
of vertebral counts for South African specimens given above are from 6 speci-
mens of Zu cristatus and 5 of Zu elongatus.
ANNALS OF THE SOUTH AFRICAN MUSEUM
34
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FAMILIES TRACHIPTERIDAE AND RADITCEPHALIDAE 35
Fitch (1964) recognized Trachipterus semiophorus Bleeker (described from
Indonesia) and T. ijimae Jordan & Snyder (from Japan) as synonyms of Zu
cristatus. All the other synonyms of Zu cristatus are based on specimens from the
Mediterranean Sea, and all the recognizable published descriptions and illustra-
tions of Zu from the Mediterranean are of Zu cristatus. It seems, therefore, that
Zu elongatus does not occur in the Mediterranean Sea.
The specimen described and figured by Whitley (1968: 45, fig. 1, as ‘Des-
modema arawatae’) is the same one reported by Hamilton (1916) from Nelson,
South Island, New Zealand as ‘Trachipterus jacksonensis Ramsay?’. This fish
appears to be Zu elongatus. The measurements given by Whitley (1968: 45), as
well as his figure, show the body depth to be 16 per cent of the standard length
(which is within the range for Zu elongatus, but not for Zu cristatus). We have
examined the specimen from French Pass, Cook Strait, reported (as Trachipterus
jacksoniensis) by Phillipps (1944) and (as Trachypterus arcticus) by McCann
(1953). Data from this specimen (NMNZ P.834) plus one other from New
Zealand (NMNZ P.2248) are not included in the diagnosis or description of Zu
elongatus given above. Counts and measurements from this New Zealand ma-
terial are as follows (P.834 given first): SL c. 425 (tip of tail separated from rest of
specimen), 315; SV 138, 108; greatest body depth 66, 48; body depth at anus
34, 22; head length 53, 43; eye diameter 15, 14; lower jaw length 31, 25. Lateral-
line scales 125, 131; pectoral-fin rays 12, 12; dorsal-fin rays, damaged, 6+132;
total vertebrae c. 88, 88; pre-anal vertebrae 39, 37; and precaudal vertebrae 32, 30.
Family Radiicephalidae
Diagnosis
Body elongate, strongly compressed, the depth decreasing gradually from
just behind head to last vertebra. Dorsal fin origin over eye. Anus well before
midpoint of body. Anal fin rudimentary, located about half-way between anus
and last vertebra. Pelvic fins rudimentary in adults, with 6—-9(?) rays in juveniles.
Lateral line continued beyond last vertebra where the canals of either side are
joined to form a large, thin-walled tube carried on the greatly elongated ‘rays’
that emanate from the ventral surface of the last three vertebrae. Length of
posterior vertebrae less than twice that of tenth vertebra. Upper jaw highly
protrusile. Skin with scattered soft papillae and numerous pores. Swim-bladder
well developed; ribs and ink-sac present.
Radticephalus elongatus Osorio, 1917
Radiicephalus elongatus Osorio, 1917: 114, figs 2-4. Walters, 1963: 456. Harrisson & Palmer,
1968: 185, figs 1-6. Palmer, 1973: 333. Parin et al., 1974: 116, fig. 13. Karrer, 1976: 179, pl. 1,
text-fig. 1. Parin, 1978: 163. Parin et al., 1978: 176.
Material
SAM-29416, c. 310 mm SL (upper jaw protracted), 420 mm TL; 70 miles
south-west of Cape Point; 600 m; Engels midwater trawl; collected by the South
African Sea Fisheries Institute, 24 May 1982.
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description (measurements in mm)
Body and tail elongate, strongly compressed. SL c. 310; body depth at
pectoral fin base 32, at anus 24,3, and at anal fin origin 11,0; body width at
pectoral base 7,5; SV c. 124; snout to anal fin c. 188; pelvic fins to vertical at rear
end of pectoral base 20,5; from pelvic fins to anus 44, from anus to anal fin 67;
head length c. 51; orbit diameter c. 13; interorbital width 8; lower jaw length 26;
maxilla width 10,3.
Dorsal fin with 160 rays, the origin above the eyes, the last ray over the
twenty-sixth preural centrum; pectoral fin rays 10 (including the short, spine-like
first ray), the longest ray 10 mm; pelvic fins damaged, most of the rays broken off
at their bases, which number 6 or 7; one 47 mm ray is still attached to the pelvic
fin base; anal fin rays 6, the longest ray 3 mm; length of anal fin base 3 mm. Gill-
rakers 3+ 8, the upper rudimentary, the anteriormost very short; branchiostegal
rays 4+3 (left side with 4+2 rays, but the membrane is torn and one ray has
probably been lost).
Lateral line begins over eye and drops to 6 mm above the anus, thence
running close to the ventral edge of the tail (1 mm above anal fin base). No
individual lateral-line scales apparent; the canal comprises a large, thin-walled
(almost transparent) tube; at about the anal fin the tube runs along the ventral
edge of the vertebral column on a shelf formed by the elongate slender haemal
spines (haemal spine of eighteenth preural vertebra 12,4 mm long; centrum
length 2,8 mm); posterior to the end of the vertebral column, the lateral-line
tubes are carried between the dorsal and ventral caudal fin ‘rays’, which are
108 mm long; width of lateral-line tube at penultimate vertebra 1,0 mm and the
vertical height of this centrum is 0,6 mm.
Total vertebrae 121, precaudal vertebrae 38, pre-anal 45; ribs are present on
all precaudal vertebrae except the first three. Length of tenth centrum (2,2 mm)
equals height of centrum at its posterior end; length of tenth preural centrum
(3,2 mm) equals 4,5 times height of centrum at its posterior end.
Upper jaw with a patch of small sharp teeth plus one much larger retrorse
tooth at each side of premaxillary symphysis; a series of 7—9 slender sharp teeth
on each side of lower jaw symphysis; 3 slender sharp teeth on vomer and 2 or 3
similar teeth on each palatine.
Body and tail silvery in life; dorsal fin base blackish.
Remarks
The specimen described above agrees well with the 597 mm SL neotype
described by Harrisson & Palmer (1968) and the 520 mm SL specimen described
by Karrer (1976). The lack of countable lateral-line scales in the SAM fish may be
due to the abrasion that the specimen has suffered and/or its smaller size. The
difference in the number of branchiostegal rays reported by the previous authors
(6 versus 7 on the right side of the SAM fish) may reflect damage to their
specimens, or perhaps the fact that the anterior two rays on the epihyal are close
together and might have been counted as a single ray.
FAMILIES TRACHIPTERIDAE AND RADIICEPHALIDAE 37
The exact nature of the caudal fin ‘rays’ is not clear; the ventral group of two
or three ‘rays’ seems to comprise the greatly elongated haemal spines of the last
two or three preural centra. The rays above and below the lateral-line tubes are
longitudinally striate, but none of these rays is segmented.
The specimen described above represents the first record of this species for
the south-eastern Atlantic Ocean. Radiicephalus elongatus is known from the
Atlantic and Pacific oceans and may also occur in the Indian Ocean.
ACKNOWLEDGEMENTS
We are indebted to Mr Alwyne Wheeler and the British Museum (Natural
History) for supplying radiographs of Trachipterus trachypterus and copies of rare
publications. Mr Wheeler also reviewed the manuscript and offered advice on the
validity of Trachipterus trachypterus (Gmelin, 1789). Dr Richard H. Rosenblatt
of the Scripps Institute of Oceanography kindly lent a specimen of Desmodema
polystictum. A radiograph of D. polystictum was supplied by the Fish Division of
the US National Museum of Natural History. We thank Dr Douglas Hoesse and
the Australian Museum for data and a radiograph of the holotype of T. jack-
sonensis. We are grateful to Dr Graham Hardy of the National Museum of New
Zealand for the loan of specimens from New Zealand. Mr Jean-Michel Vinson
did the drawings (Figs 2 and 8). Mr Len Jones contributed his observations of a
live specimen of Zu. Prof. M. N. Bruton, Miss A. E. Louw and Dr P. A. Hulley
offered constructive comment on the manuscript. Mr R. E. Stobbs produced
several excellent radiographs.
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Risso, A. 1840. Note sur une nouvelle espéce du genre Gymnetrus. Arch. Naturgesch. 6: 13-16.
RosEnBLaTT, R. H. & Butter, J. L. 1977. The ribbonfish genus Desmodema, with the descrip-
tion of a new species (Pisces, Trachipteridae). Fish. Bull. Calif. 75: 843-855.
SmiTH, J. L. B. 1949a. Forty-two fishes new to S. Africa, with notes on others. Ann. Mag. nat.
Hist. (12) 2: 97-111.
SmiTH, J. L. B. 1949b. The sea fishes of southern Africa. Cape Town: Central News Agency.
SmitH, J. L. B. 1953. The sea fishes of southern Africa. 2nd ed. Cape Town: Central News
Agency.
SmiTH, J. L. B. 1956. A new dealfish from South Africa. Ann. Mag. nat. Hist. (12) 9: 449-452.
SmiTH, J. L. B. 1965. New records and descriptions of fishes from South West Africa. Occ. Pap.
Dep. Ichthyol. Rhodes Univ. 3: 13-23.
SmiTH, J. L. B. 1966. Interesting fishes from South Africa. Occ. Pap. Dep. Ichthyol. Rhodes
Univ. 8: 83-94.
TANAKA, S. 1915. Figures and descriptions of the fishes of Japan, including Riukiu Islands, Bonin
Islands, Formosa, Kurile Islands, Korea and Southern Sakhalin. 2nd ed. Tokyo.
TorToNESE, E. 1958. Cattura di Trachypterus cristatus Bon. e note sui Trachypteridae del mare
Ligure. Doriana 2: 1-5.
TorTONESE, E. 1970. Osteichthyes (Pesci ossei), Parte Prima. Fauna Ital. 10: 1-565. Bologna:
Calderini.
WaLtTERS, V. 1963. The trachipterid integument and an hypothesis on its hydrodynamic function.
Copeia 1963: 260-270.
Watters, V. & Fitcu, J. E. 1960. The families and genera of the lampridiform (Allotriognath)
suborder Trachipteroidei. Calif. Fish Game 46: 441-451.
WEBER, M. & BEAuFortT, L. F. DE. 1929. The fishes of the Indo-Australian archipelago.
Anacanthini . . . Heterosomata, Berycomorphi, Percomorphi (Kuhliidae . . . Centropomi-
dae). Leiden: E. J. Brill.
WuitLey, G. 1927. Studies in Ichthyology. No. 1. Rec. Aust. Mus. 15: 289-304.
WuitLey, G. 1968. A check-list of the fishes recorded from the New Zealand region. Aust. Zool.
15: 1-102.
ue
~
|
cb
Cray,
6. SYSTEMATIC papers must conform to the Jnternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
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not acceptable.
In describing new’species, One specimen must be designated as the holotype; other speci-
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not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. ‘Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
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Biological Abstracts.
P. C. HEEMSTRA & S. X. KANNEMEYER
THE FAMILIES TRACHIPTERIDAE AND
RADITCEPHALIDAE (PISCES, LAMPRIFORMES)
AND A NEW SPECIES OF ZU
FROM SOUTH AFRICA
VOLUME 94 PART 3 JULY 1984 ISSN. 0303-2515
CAPE TOWN
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Examples (note capitalization and punctuation)
BULLOUGH, W..S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIscHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FiscHer, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270.
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 94 Band
July 1984 Julie
Part 3 Deel
LOBEDU MATERIAL CULTURE
A COMPARATIVE STUDY
OF THE 1930s AND THE 1970s
By
PATRICIA DAVISON
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
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Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap
LOBEDU MATERIAL CULTURE
A COMPARATIVE STUDY OF THE 1930s AND THE 1970s
By
PATRICIA DAVISON
South African Museum, Cape Town
(With 119 figures)
[MS accepted 1 November 1983]
ABSTRACT
In this museum and field study a collection made by E. J. Krige and J. D. Krige between
1936 and 1938 is described and compared with data recorded 40 years later by the author.
Within the descriptive framework, material is ordered according to three categories of re-
sources—natural, domestic, and imported. Dependence on the indigenous environment is
shown to have been important in the traditional technology and in understanding the value with
which certain materials were invested. Effects on the material culture of the transition from a
subsistence to a money economy are discussed. It is noted that women have proved both con-
servative and adaptable in changing economic circumstances and that the practice of certain
crafts has increased to supply both domestic needs and an informal market. Conclusions are
drawn regarding the interrelated nature of material and social change.
CONTENTS
PAGE
MEE HO GU CLIO Me hoo ties sek a asd ae Sie eae ee 42
RIS CORIC AOU EMC = oe al aia. euskal eee ea ey age Nea 44
ESV OMIMETID eo hi Se a Sane en are ee aD Pea ee 47
| DYSSgaVOVEd C2) 0) 0 het ee eR san cia tric. cea cinnics cigs ois 49
Socialicontexts ca ay ieta sala Oh UE ee ee Sik
MC Slt OLO ON e eee. aee vce acacyes ony ie eaets ee aha serene eae 53
Wyse ofmaturalsreSOUrCES:., S45. Sl. See ne ee eo es 54
Uy ire ee Ne costed «cute hens tea eee Os a 54
Grasses andiother plant fibies 7. sees nee eee 68
WOO Gran Gime eG ce sone cose eee ee eee A pine Saar 100
SSO TUE eae ane a os A Se ee rt te le 143
Summianysand commento! ss 4 ee se oe eer eee 147
Wsetofidomestic RESOUICES).... =. asi eee ee ee eee 150
GOUndSS Ake Reta oe Jal ee eto he ae et ee oe ane en 150
Skansiandehrdesy 2.200 0. a0. aac See oe ne oe ne See 156
AOR AN GD OMNEH. Ae Skok ee ee a AT Orne Huet ra oor ae 164
IB) RIA Sey 8 ee 8 iit ees cil tea eS eh a ates ea 169
Summanysand comment (ec. yes ee ese oe ee 170
WseotimpontedimatertalS a4. shee acess Soe ee 7A
INCRE AGAR ar oa ef ere rot OO OME tence RACERS. | EN 171
Glassiamidislee Ula ek ee) e n a rR ea 178
COTS Yd ne See aa eH ee a a eT Pee eee ee YA 183
Sa ere a me A ee Be 2 als CORO ate TEE 185
Summany-andicomment 4.25 on shaae eas soem ee 187
41
Ann. S. Afr. Mus. 94 (3), 1984: 41-201, 119 figs.
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
PAGE
Material’cultuneandisocialichangen s.70 oa eee 189
Historical, environmental and social change c. 1880-1980 ... 189
Change imimatenaliculturec, 1940=S80 9 ee ee 192
Conclusions regarding the nature of change ................ U7)
ACKNOWIEGSEMIEIIS Was an yu 5 Oe, Gatch ae ancy EI Cue ae ae 199
IRCIEREHCES nto eS cee Moe ee ne HAE Cee Che ee ares 199
INTRODUCTION
The Lobedu of the northern Transvaal have been the subject of anthropolo-
gical study from the 1930s to the present. Their social structure has been de-
scribed in detail (Krige 1931, 1932, 1938, 1964, 1974, 1975a, 1975b, 1975c, 1981;
Krige & Krige 1943, 1954; Monnig 1961, 1963) and the ethnography has pro-
vided the basis for comparative analytical studies (Leach 1961; Sansom 1974;
Kuper 1975, 1982). Thus far little has been published on their material culture.
The aim of this paper is to describe traditional Lobedu material culture as rep-
resented in a collection (hereafter referred to as the Krige Collection) made be-
tween 1936 and 1938 by social anthropologists E. J. and J. D. Krige, and to
record changes observed 40 years later during fieldwork undertaken by the
author. .
The Krige Collection was made with the purpose of bringing together ob-
jects that represented the full range of the material culture of a single society at
a particular time. It was built up during the course of anthropological research
involving participant observation among the Lobedu people over a continuous
period of nearly 18 months. In 1970 the collection, together with short notes on
each item, was presented to the South African Museum, Cape Town, where it is
now housed. It is primarily on the Krige Collection and notes that the descrip-
tion of Lobedu technology in the 1930s is based.
The collection comprises 248 specimens (SAM-—9671 to 9919) and represents
almost all aspects of Lobedu material culture with the exception of very large
objects and immovable structures. The number of examples of each type of ob-
ject, however, is not large enough to assess the range of variability. The area in
which the collection was made covered most districts of the original Modjadji
Location demarcated in 1892, as well as farms and Crown Land to the north and
the area of the old Medingen mission station. For the purpose of this study, it is
assumed that the artefacts in the collection are representative of the material
culture in the 1930s unless the collector’s notes state otherwise. The impression
gained from the collection is essentially qualitative but the large body of related
data allows the material culture to be viewed within wider social, economic, and
environmental perspectives. The anthropological literature on the Lobedu pro-
vides valuable contextual data for the collection. In addition, E. J. Krige has
made available unpublished photographs taken during the 1930s, and relevant
unpublished data relating to her fieldwork in the 1930s and during the period
1962-1980, and has further assisted the study by providing additional informa-
tion on the situation in the 1930s (Krige 1982: 1-45), which allowed a more
i) es
LOBEDU MATERIAL CULTURE 43
accurate comparison to be made with the 1970s data. Unacknowledged photo-
graphs in the text were taken by the author.
Field research undertaken in September 1973, April 1975, June 1976, and
April 1978 was planned with the aims of recording in detail the technological
processes used by the Lobedu and of comparing their recent material culture
with that represented in the Krige Collection. A total of 3: months was spent in
the research area. It was considered important to collect data at different times
of year in order to observe the productive activities associated with the seasonal
cycle. As far as possible this was done, with the exception of the summer wet
season. While doing fieldwork it was necessary to commute daily between the
Lobedu reserve and Duiwelskloof, some 20 km away, as permission to camp in
the reserve was not granted by the local magistrate. With the guidance of
Simeon Modjadji, a member of the royal family living at the capital and one of
the assistants of Krige & Krige, and by using the Krige Collection as a baseline
record of the material culture, more was achieved in a relatively short period of
fieldwork than would otherwise have been possible. Charts with the vernacular
terms and photographs of all the items in the Krige Collection were used to
record whether traditional artefacts were still currently in use. Frequency of
occurrence was noted and absence of any item was followed up with inquiry
regarding replacement by alternatives and reasons for disuse. The focus of this
investigation was Modyjadji’s village, the capital, which has been continuously
occupied for the past 70 years, as well as smaller and newer villages on the sur-
rounding hills and in the Molototsi valley. Areas beyond the boundaries of the
old Modjadji Location, the Medingen mission area and Gakgapane were not in-
cluded.
Once it had been established that many of the traditional crafts, notably
pottery, basketry and wood-working, were still active, the processes involved
were observed and recorded during the normal work routine of the craftsmen.
This was supplemented by demonstrations of particular techniques no longer in
regular use but still remembered. Where possible all stages in the processes were
photographed in situ and examples of partly made artefacts collected for further
study. Specimens of raw materials were collected, and the source and availabil-
ity noted; preparation of materials, tools used, and techniques employed were
observed in action and confirmed in interviews with the craftsmen. Inquiry was
made into specialization and the transmission of skills from generation to gener-
ation.
In the case of processes that were no longer practised, information was
sought from older people who may have recalled techniques formerly used. No
satisfactory information on iron-smelting was obtained as the complex tech-
nology involved seemed to have been forgotten even by the older generation.
Accounts of skin-working technology were obtained mainly from people who
had had past experience but no longer practised it.
Specimens from Modjadji’s area in the private collections of M. Klapwijk
(now part of the Transvaal Provincial Museum Service collections housed at
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
Eiland in the northern Transvaal) and J. Witt, both of Tzaneen, and in the col-
lection of the National Open Air and Cultural History Museum in Pretoria were
recorded and photographed to supplement the Krige Collection.
The descriptive framework adopted orders the data according to the ma-
terials utilized in the technology. The concept underlying this approach is that
there is an interaction between the resources of the environment, the technical
skills used in exploiting these resources, and the social context in which the tech-
nology operates. Following this framework, objects made of different materials
and the related processes are described in separate sections. In reality, however,
everyday activities integrate artefacts made of clay, wood, grass, gourds, and a
variety of other materials. Cross-references in the text indicate where objects
are used concurrently and where similar processes are applied to different ma-
terials. In conclusion the separate sections are drawn together in a discussion of
change in the material culture.
Simeon Modyjadji acted as interpreter throughout the study. There is no
standard orthography for the Lobedu dialect, which has characteristics of both
Venda and North Sotho. North Sotho is the language taught in schools. In the
ethnographic literature spelling is inconsistent. For example, the bi-labial v in
Lobedu is inconsistently rendered as b or v. The spelling ‘Lobedw’ has been used
as this is the standard rendering in North Sotho (E. O. J. Westphal 1982 pers.
comm.). The vernacular terms have been given in khelovhedu dialect, spelt by
using Venda for the consonants and Sotho for vowels following the advice of
N. J. van Warmelo (1977 pers. comm.).
HISTORICAL OUTLINE
Oral traditions recorded by Krige & Krige (1943: 1-12) trace Lobedu his-
tory to the disintegration of the Karanga state north of the Limpopo River dur-
ing the sixteenth century. The Lobedu royal line is thought to descend from one
of the three great dynasties, Mwene Mutapa, Torwa, and Changamire. In the
late sixteenth or early seventeenth century a break-away group migrated south-
ward across the Limpopo (Alpers 1970: 217) taking with them the knowledge of
rain-making and the sacred glass beads that were among the insignia of
chieftainship. The early Lobedu moved into the Lowveld and established them-
selves as rulers in their present environment by subduing the previous inhabi-
tants, thought to have been scattered groups of Sotho-speaking people. A
period of internecine strife was followed in about 1800 by the accession of the
first woman chief, Modjadji I, who restored political stability and achieved wide-
spread fame and power as a rain-maker.
Lobedu oral traditions have parallels among those of the Venda who
crossed the Limpopo later in the seventeenth century and also trace their ances-
try to the Karanga (Stayt 1931: 12). During the post-migratory period bonds be-
tween Lobedu and Venda were strengthened through trade, marriage alliances
and the Venda practice of coming to Modjadji to supplicate for rain.
LOBEDU MATERIAL CULTURE 45
Notable similarities in language, custom and material culture confirm their past
association.
During the nineteenth century Modjadji’s influence spread to other Sotho-
speaking people living in the Lowveld, including the Letswalo, Kgaga, Thabina,
and Narene. Culturally the Lobedu have much in common with these neigh-
bouring people, as do they with the Phalaborwa people further east (Du Toit
1968; Hammond-Tooke 1981; J. D. Krige 1937). By comparison, they have little
in common with Sotho-speaking people living in the different environment of
the Highveld (Krige & Krige 1943: 306; Hammond-Tooke 1981: 154).
From about 1840 onward successive waves of Tsonga-speaking refugees en-
tered the Lowveld as a result of the social disruption caused between 1835 and
1840 by the invasions of the Ndwandwe warrior Shoshangane in what is now
southern Mozambique, and after 1856 by the wars of succession followed by at-
tenuated raiding, drought, and related famine and disease. Prior to this time a
few Tsonga-speaking groups had been assimilated as Modjadji’s subjects but the
large number of nineteenth-century immigrants who had to settle in the un-
healthy low-lying areas were not absorbed through intermarriage and were re-
garded as outsiders. This exclusion was reflected in the refusal of Modjadji to
allow implements of Tsonga manufacture or origin, such as wooden mortars for
stamping maize and the tambourines used in the maldb6 cult, to be adopted at
the capital (Krige 1931: 233; Mantwa Modjadji 1981 pers. comm.; Krige 1982:
21).
During the second half of the nineteenth century expansion of the Zuid-
Afrikaansche Republiek into Modjadji’s territory initiated a period of conflict
and fundamental change. From as early as 1855 the Volksraad imposed on chiefs
in the Soutpansberg area an annual tax of five head of cattle (or five elephant
tusks, or twenty-five copper rods, or twenty leopard skins) and appointed offi-
cials to exact this tax (Grimsehl 1955: 209). In 1861 Albasini, the official respon-
sible for the Lobedu area, had cause to complain that Modjadji refused to pay
the annual tax. Later that year a commando force took action against her and
confiscated a large number of cattle and small livestock as penalty. In the years
that followed Modjadji remained intransigent and prepared to resist further in-
vasions of her territory (Grimsehl 1955: 205).
In 1866 the Volksraad changed the law regarding taxation so that Africans
could be taxed individually and in 1876 there was a further revision according to
which every male over 20 years of age was required to pay 10 shillings per year,
and, in addition, a hut tax of 10 shillings per year was payable. By the same rul-
ing it was decided that ‘locations’ would be assigned to the more important
chiefs who would be paid a salary and be required to keep law and order, and
supervise the collection of taxes (Grimsehl 1955: 206). Modjadji, however, was
not prepared to co-operate and her indignation was further aroused when in
1886, before the ‘location’ boundaries had been fixed, the Volksraad placed ‘oc-
cupation farmers’ on land already occupied by her subjects. The Lobedu resisted
with arms and by burning down farmers’ houses and raiding cattle. In 1890 a
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
commando force was sent to Modjadji to exact taxes, confiscate arms, and settle
the unrest in the area (for details see Grimsehl 1955: 211-221).
In 1892 the boundaries of the Modjadji Location were fixed despite strong
opposition from Modjadji’s councillors and headmen. The area was less than
one tenth of the area that Modjadji had previously controlled. Discontent over
this dispossession of land continued and eventually led to armed confrontation
in 1894 when General Joubert finally crushed Lobedu resistance and confiscated
about 10 000 head of cattle in retribution (Grimsehl 1955: 236-250).
During these last unsettled decades of the nineteenth century the Berlin
Missionary Society established a station in Modjadji’s area (Reuter 1905: 249).
In 1881 Reuter founded the mission settlement at Medingen and started prosely-
tizing among the Lobedu but with little early success. The drought of 1881-2
was attributed to the presence of the mission and Khashane, a Lobedu evangel-
ist who was treated with great suspicion, was eventually murdered in 1884.
Reuter, however, persevered in his cause, established a school, trained young
men in technical skills, and interceded on behalf of the Lobedu in negotiations
with the Volksraad over their territorial boundaries.
In 1895 Modjadji II followed the tradition of her ancestors in committing
ritual suicide. By the end of her reign her territory had been greatly reduced and
the value system that sanctioned her authority was under pressure.
In the years that followed white encroachment continued. The appropria-
tion of resources undermined local production, as did the importation of mer-
cantile goods. Higher taxes payable in cash and an increasing need for money to
buy commodities forced men to become migrant labourers. By 1920 a railway
line linked the Lowveld to the industrial centres of the Transvaal and facilitated
mobility. Pass Laws, however, which had been introduced as early as 1901, re-
stricted the movement of migrants.
One of the results of the South African War was that many Africans moved
back on to the land that had previously been appropriated. None the less, press-
ure on arable land was high. In 1910 it was reported that ‘In the vicinity of Dui-
welskloof and around Modjadji’s location some farms are so densely populated
with squatters that it is difficult for them to find sufficient land to cultivate with-
out interfering with white settlers’ (Blue Book on Native Affairs 1910: 269). The
introduction of the plough, which allowed maize to be more effectively culti-
vated, only partly offset the pressure on arable land as in many cases, owing to
shortage of draught animals, the plough was not set deep enough to break more
than the top-soil. Furthermore, use of the plough is dependent on good rainfall
and suitable topography, with the result that in dry seasons and in steep areas
hoe-cultivation continued.
Regulations for the dipping of cattle were received with suspicion by the
Lobedu, wary of interference. Public-health measures were introduced, quinine
was freely distributed but malaria remained prevalent in the low-lying areas, es-
pecially in the wet season. The annual recurrence of malaria in the Lowveld was
one of the factors that discouraged white economic development of this area
LOBEDU MATERIAL CULTURE 47
until much later in the 1950s after the large-scale use of pesticides had lowered
the risk of disease. During the 1930s when Krige & Krige carried out their field-
work, the Lobedu area was still relatively isolated and despite external con-
straints a subsistence economy was still viable.
During the years between the mid-1930s and the mid-1970s, which are the
main focus of this study, the Lowveld was opened up to far more intensive white
economic exploitation as a result of the eradication of malaria and improved ac-
cess with the construction of new roads. New employment opportunities were
provided for blacks living in the area and more money came into the local econ-
omies. At the same time population of the rural areas increased following the
implementation of influx control in urban centres. The resultant pressure on
arable land made subsistence agriculture an impossibility for many living in the
Lobedu area and the economy became increasingly monetized.
Also during this period political constraints increased. The Bantu Author-
ities Act of 1951, which provided for the establishment of tribal, regional and
territorial authorities, laid the foundation for the later ‘homeland’ policy. Within
this system the authority of the traditional chiefs, councillors and headmen is
greatly circumscribed by the imposed political structure and appointed officials.
The Lobedu district was included in the North Sotho homeland, Lebowa, which
in 1972 was declared a self-governing area within the Republic of South Africa.
The challenge to traditional values initiated during the late nineteenth cen-
tury gained in intensity during the twentieth century as the authority of the
chiefs was undermined, as education facilities improved and mobility increased.
In 1959 Modjadji III died naturally, having rejected the rite of suicide, and was
succeeded by her daughter, Modjadji IV, who died in 1980 and was in turn suc-
ceeded by her daughter. Modjadji V is represented on the Lebowa Legislative
Assembly by her brother, Legugela.
ENVIRONMENT
Modjadji’s territory is situated in the foothills of the northern extension of
the Drakensberg in the eastern Transvaal, at less than 1 000 m above sea level.
The topography varies from relatively high mountainous areas to low-lying river
valleys and plains. Climatic conditions differ considerably with topographic
variation and, despite mountains and rivers, there are stretches of land with
little or no water. In general the lowlands are less well watered and hotter than
the highlands where the temperatures are seldom extreme. The predominant
type of soil is a fertile reddish loam suitable for raising a variety of grain crops.
Much of the highland area is too steep for ploughing (Hattingh 1975: 60) but
well suited to terraced hoe-cultivation. Clays of different types are found in
many places, mainly in river-banks, and are used extensively in the technology.
The area falls within the summer-rainfall region. The rains normally start in
October and continue to March. On average January and February are the wet-
test months, having falls of about 150 mm. Rain is usually plentiful on the
mountains but less predictable on the plains; in one season there may be good
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
rains in one area and drought in another. This unpredictability of rainfall was a
major source of insecurity in the agricultural economy. By having fields in differ-
ent areas and by planting drought-resistant sorghum as well as maize the risk of
total crop failure was reduced. In times of drought wild foods formed an import-
ant component of the diet.
The natural vegetation is bushveld savannah rich in trees, shrubs, creepers,
and grasses. Most of the area is of the veld type described by Acocks (1975:
27—28) as ‘Lowveld sour bushveld’, which provides poor-quality grazing. The
hillslopes are thickly wooded including, in one place, a unique forest of cycads.
A nineteenth-century traveller (Nelson 1973: 221) described Modjadji’s area as
‘some of the richest, wildest and most romantic scenery to be met with .. .’.
The valleys have open woodland vegetation with an abundance of large trees,
among which the marula and a number of Acacia, Combretum and Ficus species
are prominent. Traditionally the Lobedu made extensive use of these plant re-
-sources both for food and in the manufacture of artefacts. The particular value
of the marula tree is clearly reflected in the material culture and in the diet.
Oral traditions also reflect the importance of natural resources. Kruger
(1936: 91) translated a vernacular account as follows: ‘Now the Lobedu were
looking for a nice country with fertile soil and with ant-hills. They sent the
young men in advance to look for such a country. And indeed . . . they found
the right country having marula-trees and ant-hills. . . .. Thus recorded are the
essential elements of Lobedu subsistence, a natural vegetation rich in useful
trees and wild fruits, edible insects, and fertile arable land.
By the 1940s natural timber resources had been seriously depleted, mainly
through the clearing of land for fields and settlement, and legislation to protect
certain indigenous trees had been introduced. Many of the trees, formerly used
by the Lobedu, were affected and could no longer be cut down without payment
of a fee. Dead-wood, however, continued to be used without restriction both for
fire-wood and for making utensils, and illegal use of wood was not uncommon.
Formerly, when the indigenous forests and woodlands were thickly in-
habited with game, hunting was an important source of meat as well as skins and
horns for domestic use. Ivory, horns and skins were also of considerable econ-
omic importance in external trade through Delagoa Bay. Modjadji’s territory lay
just south of the rich Soutpansberg hunting-grounds known to have been ex-
ploited from the early eighteenth century by Venda and Hlengwe hunters (Wag-
ner 1980: 324). By the mid-nineteenth century the introduction of fire-arms by
white hunters had changed both the nature of hunting and the scale of exploita-
tion.
Modjadji’s territory was undoubtedly rich in game but, according to Krige
(1983 pers. comm.), although the Lobedu hunted for meat, they did not play an
active role in hunting for the ivory trade as did their Venda and Tsonga neigh-
bours. In the mid-nineteenth century ivory was a common medium of exchange
and Modjadji received ivory in tribute from hunters in her territory and from
neighbouring chiefs such as Mawewe (Das Neves 1879: 104). Tribute brought to
LOBEDU MATERIAL CULTURE 49
Modjadji from supplicants for rain ensured that she benefited from the game
trade while not actively participating. In the Lobedu context, tribute in goods
had rain as its reciprocal counterpart. By the 1930s not only had the game popu-
lation been drastically reduced but both hunting and the possession of fire-arms
were restricted by law.
The Lowveld was subject to infestation by a number of insects that threat-
ened both human and animal life. Malaria was prevalent in the low-lying areas,
especially during the wet seasons. Tsetse fly (Glossina sp.) was widespread and a
serious threat to cattle, which were also subject to epidemics such as east-coast
fever and rinderpest. Swarms of locusts threatened crops and wild plant foods;
wood-boring termites attacked structures and utensils, and grain stores had to be
protected against weevils. Not all insects, however, were regarded as pests.
Edible termites (Macrotermes sp.) formed an important part of the diet, which
was also supplemented by grasshoppers.
There are some indications that prior to the alienation of grazing-land and
the rinderpest, herds were relatively large, despite the presence of tsetse fly.
Reuter, reporting to the Holloway Economic Commission in 1930, commented
that when he came to the area in the 1880s ‘the lands were lying idle but the
country was full of cattle’. Not only were cattle more numerous but they were in
better condition because grazing-land was plentiful. It was the rinderpest of
1896, together with lung-sickness and east-coast fever, that destroyed almost all
the cattle in the area. Recovery thereafter was slow and restricted by limitations
on grazing-land and the poor quality of the sourveld pasture. The small number
of approximately 3 000 cattle recorded in the Lobedu reserve in 1936 (Krige
1941: 2) seems to be less related to the effects of tsetse than to the disruption of
grazing-patterns brought about by the fixing of territorial boundaries.
DEMOGRAPHY
The Lobedu are a heterogeneous group of originally unrelated people who
over the centuries have become linked together through a network of social
relations and who now tend to conform to a common cultural pattern. The
Lobedu who trace descent from the original founding group comprise less than
10 per cent of the Lobedu as a whole (Krige & Krige 1943: 85).
Before the territorial boundaries were drawn up in 1892, Modyjadji was ac-
knowledged as chief over the vast area bounded by the Great and Little Letaba
rivers (Krige & Krige 1943: 13). The demarcation of the Modjadji Location
(Fig. 1) in 1892 limited Modjadji’s territory to approximately 17 500 morgen
(179 km?) despite complaints by Lobedu headmen to the Location Commission
that this was insufficient land to support their people (Grimsehl 1955: 228-231).
During the 1930s the Lobedu extended their land to approximately 389 km? by
the purchase of farms to the north and east (Krige & Krige 1956: 12), and in the
1960s trust lands to the north were included under Lobedu jurisdiction.
In 1904—5 (Report of the Transvaal Native Administration) the population
of the original Modjadji Location was 14 434, giving a mean density of
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
MODJADII
fo OES
a
Duiwelskloof
30°15’ 30° 30'
Fig. 1. Original Modjadji Location demarcated in 1892 and BoLobedu magisterial district,
1979:
81 persons/km?. In 1936 the population of the extended territory was estimated
at 33 000 (Krige & Krige 1943: 13) giving a mean density of 85 persons/km?. By
1980, according to the official census for the BoLobedu district, the population
had risen to 116 643, and the average density was well over 200 persons/km?. In
areas of most concentrated settlement the density is considerably higher. The
resultant shortage of arable land has led to an inevitable move from agriculture
to dependence on wage labour.
In the late 1970s over 65 per cent of males were migrant labourers and ab-
sent from the reserve at any one time, compared to 37 per cent in the mid-1930s
(Krige 1981: 151). The 1970 census figures indicate that women outnumber men
LOBEDU MATERIAL CULTURE Sit
in the ratio 3:2, a proportion effectively increased by the absence of men work-
ing outside the district. Relatively few Lobedu women are employed as migrant
labourers but there is a trend for women to work as daily labourers on farms,
tea plantations, and in light industry (Krige 1981: 155).
The traditional settlement pattern was one of dispersed villages situated
mainly on the highlands and hillslopes. A village was composed primarily of an
extended family, the size of which varied considerably with the number of wives,
children, and other dependants. The largest settlement was, and still is, the capi-
tal, mosatha (see Fig. 73A), which in 1975 had a population of over 500 people
living in a concentrated clustering of dwelling-units around the central court-
yard. Most settlements are much smaller, consisting of between two and ten
groups of huts. In the 1930s there were a few larger villages of up to eighty huts
(Krige & Krige 1943: 17). New homesteads tend to be very small, sometimes
having only two huts, and they are sited wherever land is available. Resettle-
ment schemes, planned for the whole area, have been implemented gradually,
causing the division of extended families. Increased mobility is reflected in the
tendency for new settlements to be sited near roads leading out of the area (Hat-
tingh 1975: 64).
SOCIAL CONTEXT
Lobedu social structure has been well documented and analysed by Krige
(Krige & Krige 1943; E. J. Krige 1964, 1975a, 1975b, 1975c, 1981) and re-
analysed by Leach (1961) as well as Kuper (1975, 1982) who suggests that the
Lobedu social order can be understood as an extreme variant of the Sotho-
Tswana system. While sharing many cultural characteristics with other Bantu-
speaking people in southern Africa, the Lobedu have certain distinctive features
or emphases that find their closest parallels among neighbouring Lowveld Sotho
people and the Venda.
Among the Lobedu it is not unusual for women to hold high public office
and, indeed, the last five rulers have been women. Women play significant roles
in the social structure both as sisters (in which capacity they are frequently
leaders in ritual) and as wives (who are important in the agricultural economy).
This importance of women in the social structure is a factor that has encouraged
conservatism and adherence to tradition.
Although descent is patrilineal and residence after marriage viri-local, there
are no corporate, property-controlling patrilineal groups. In practice matrilateral
kin are as important as patrilineal kin. This is best understood with reference to
the preferred form of marriage which is between a man and his mother’s
brother’s daughter, his matrilateral cross-cousin. After a number of generations
this marriage pattern results in a merging of kin groups and an emphasis on bi-
lateral kinship.
An intricate network of royal marriage alliances has operated through time
to extend and consolidate Modjadji’s control over a wide area. From throughout
her area of influence men sent daughters to Modjadji as a form of tribute or to
Sy ANNALS OF THE SOUTH AFRICAN MUSEUM
solicit aid. These girls were accepted as ‘wives’, vhatanoni, and were expected to
remain chaste and to serve Modjadji as they would a husband. Those who did
not remain chaste were given in marriage to headmen or to royal kinsmen, or
they could be ‘placed’ to rule over a district. The network of affinal links so
formed, and implying both rights and obligations, has been an important inte-
grating force within the social and political structure (Krige 1975b: 63-64).
When Modjadji ‘placed’ a wife, no bride-wealth was exchanged but it was ex-
pected that in time a daughter would be sent to the Queen, so ensuring continu-
ity of the affinal link between ruler and subject.
It has been noted that the role of a sister was central to the Lobedu social
system (Krige & Krige 1943: 70). The importance of the sister—brother bond
was related to the pattern of marriage. Before money was used as bride-wealth,
one of the few ways in which a young man could acquire the cattle to contract a
‘marriage was to use his sister’s bride-wealth. As most cattle were tied up in mar-
riage transactions, ideally every brother had a sister whose bride-wealth he
could use in order to marry and establish a ‘house’. This ‘cattle-linked’ sister had
certain rights in the ‘house’ which she had helped to establish. In particular she
had a claim to a daughter of that ‘house’ as a wife for her son, 1.e. as a daughter-in-
law who would cook for her and help her with other domestic tasks, especially
in her old age. This would, in fact, be the preferred cross-cousin marriage,
and the girl would become the chief wife and bear the heir. Should a ‘cattle-
linked’ sister not have a son, she still had a right to a daughter of the ‘house’
which she had helped to found. In this case the sister might ‘marry’ the girl her-
self, using the bride-wealth of one of her own daughters, and appoint a man to
father children in the name of the son that she lacked.
The influence of a sister was often further reinforced by her position as
ritual leader in her brother’s household. In this capacity she would officiate at all
important ceremonies and would be the most appropriate person to intercede
with the ancestors on their behalf.
Young wives (with the exception of vhatanoni) had lower status than sisters.
On marriage the rights to a woman’s children and to her labour were transferred
from her own family to that of her husband. At first a young wife was subordi-
nate to her husband’s mother, cooking at her hearth and using her utensils.
Later, often after the birth of a child or when a younger wife joined the house-
hold, she was given her own mosha or ‘house’. In a polygynous family each wife
and her children formed an independent economic unit.
The subsistence economy was based on agriculture. Three seasons were
recognized: khelemo the time of hoeing (September—January); /etavula, sum-
mer, when the foods were plentiful (February—May) and maria, winter, when
the fields were bare (June-August). Grain provided the staple diet, supple-
mented by vegetable produce and veld foods. Neither milk nor meat was a
major component in the diet.
Division of labour was not rigidly defined. Both men and women worked in
the fields to produce the essential crops of maize, sorghum, and millet. Women
LOBEDU MATERIAL CULTURE 53
also cultivated fields of ground-nuts, beans, gourds, and other vegetable pro-
duce. Children helped with weeding and in protecting the crops from predators.
During the dry winter months both men and women worked on renewing the
homestead and its material contents. As migrant labour increased women took
over many of the tasks formerly done by men thus ensuring some degree of
stability in the productive economy.
Religion centred around belief in the ancestors. A creator, Khuzwane, was
acknowledged but was thought to be remote from everyday life. The ancestors,
on the contrary, were thought to be intimately involved with the daily lives and
fortunes of their descendants. Their displeasure could cause misfortune, ill-
health and, in the extreme, drought. To ensure health, fertility, and prosperity,
it was necessary to approach the ancestors by making offerings of beer or grain
at their shrines or by invoking them through sacred thugula objects, such as heir-
loom beads or hoe-heads. The presence of ancestor shrines, which often took
the form of a small mound of clay in which thugula objects were embedded, is
an aspect of Lobedu religion also found among other people of the Lowveld and
is distinctive to this area (Hammond-Tooke 1981: 153).
Cultural parallels with other Lowveld people are also reflected in the im-
portance of drums, which are sounded on almost all ceremonial occasions, and
in the vyali and vuhwera rituals which are essentially fertility rites, as well as
complex initiation ceremonies (Krige & Krige 1943: 126-140). Four sacred
drums are believed to be mystically associated with the Queen and the welfare
of the people. Formerly they were beaten at the g6mana initiation to appeal for
rain and in thanksgiving for the harvest, practices that have long since fallen into
disuse.
Modjadji was in the powerful position of being able to intercede with the
royal ancestors on behalf of all her subjects. Her ritual power as a rain-maker
was a vital factor in validating the chieftainship. Praised as “Transformer of
Clouds’, she was believed able to make or withhold rain, the life-giving force on
which fertility was dependent. The whole seasonal cycle was regulated by Mod-
jadji, not only through rain-making but in the tasting of the first fruits and the
ritual ‘biting’ of the first termites.
Rain was symbolic of order in nature. By association with rain, cool or
damp places and the materials found there were imbued with positive forces. A
river stone buried at the entrance to a village was thought to ‘cool’ or counteract
the forces of evil. Coolness denoted a state of harmony, conversely ‘heat’ was
perceived as a force leading to disorder (Krige & Krige 1954: 68). These two
concepts are important in understanding the symbolic content of the material
culture and the Lobedu perception of their environment.
TECHNOLOGY
Technology has been simply defined as the complex of learnt behaviour that
gives rise to material culture (Spier 1973: 1). It is behaviour concerned with
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
making things—the materials, tools, skills and processes employed to fulfil ma-
terial needs. It is also adaptive behaviour, the means by which man interacts
with and modifies his environment.
Adaptive strategies, however, do not necessarily correlate directly with
available resources. The environment provides a range of resources but cultural
responses account for selectivity, for methods of exploitation, and for the value
with which certain resources are invested. Ideological as well as economic fac-
tors influence the manufacture and use of artefacts.
In the framework for description adopted here, objects are grouped accord-
ing to the materials of which they are made and three categories of resources are
reflected: natural resources indigenous to the environment, domestic resources
that are cultivated or the product of animal husbandry, and imported materials.
Within this arrangement it is unavoidable that artefacts used concurrently but
made of different materials are described separately. Context and related use,
however, are indicated and illustrated in the photographs.
USE OF NATURAL RESOURCES
CLAY
Sources of clay are well distributed in the Lobedu area. The weathering of
granitized rocks, which predominate in the area, yields clay minerals and quartz.
Through selective transport of these materials by water or wind erosion the
lighter clay minerals may be concentrated locally, as in the case of clay beds in
river-banks. This is the source most frequently utilized by potters. If decomposi-
tion of the rocks takes place on site with no transport, the clay will be mixed
with quartz particles as the latter are virtually inert and resistant to decay. In
this case the clay requires sorting and sifting before use in making pottery.
Muds of rougher texture found generally throughout the area are used for
plastering walls and floors of huts, for building courtyard walls and ledges and
for making sun-dried bricks (see p. 138). Natural earth pigments such as ochres
and chalks occurring in pockets in certain places, are collected, ground into a
fine powder and used for colouring walls (Fig. 2), clay utensils and, in certain
circumstances, the body. Graphite, which occurs less generally in the area, is
also used as a colourant.
Pottery techniques
The manufacture of pottery, described in an earlier paper (Davison &
Hosford 1978), is summarized below.
Among the Lobedu, pottery is always made by women, some of whom are
specialists and make pots for sale as well as for their own use. Most potters learn
how to make pots by watching and following the technique of an experienced
potter. The craft is not necessarily passed on from mother to daughter and the
present generation of young women show little interest in learning the craft.
In general, pottery manufacture can be separated into at least five stages:
digging the clay, preparing it, forming the pot, drying it, and firing. Decorating
LOBEDU MATERIAL CULTURE 55
Fig. 2. Ochre decorations being applied to wall of courtyard, Modjadji’s village, 1973.
>
a 22 y
erty
3 ee
is ieee
Fig. 3. Collecting clay from river-bank, Molototsi valley, 1975.
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
or burnishing often occurs during the drying stage and may be retouched after
firing.
Lobedu potters in the vicinity of the royal village collect clay, ledzoba, from
two deposits, one about a kilometre from the village, the other about three kilo-
metres away. Both clays, mainly kaolinite, are dug from the banks of dry river-
beds (Fig. 3), but the clay from the further source is darker and finer than the
yellowish, coarse clay of the nearer site and needs less preparation for use. The
finer clay requires only kneading and the removal of extraneous matter; the
coarser clay is dried, crushed, and ground to a fine powder between grinding-
stones, covered with water and allowed to stand for a few days before being
worked into a homogeneous mass by being pounded with a wooden pestle on a
concave stone (Fig. 4A). No tempering material or filler is added to alter the
plasticity of the clay.
All Lobedu pots are formed in a similar way. The start is made by placing a
rough ring of clay on a movable support (a potsherd, wooden dish, enamel basin
or plate) (Fig. 4B). From the ring the lower part of the pot is formed by pinch-
ing, scraping and smoothing by hand, adding pieces of clay (Fig. 4C—D) and,
when it is roughly the required shape, by smoothing it with the shell of a seed-
pod from the mothema creeper (Bauhinia kirkii). The pod, thema, is left in
water to become pliable and is then used in a variety of ways, the convex side
for smoothing, -shevola, the concave side for scraping, and the edge for shaping.
At first the thema is used to smooth the outer wall in an upward, diagonal move-
ment while the other hand supports the wall on the inside and presses lightly to
heighten and shape the sides. Small rolls of clay are added to the sides and
worked into the body of the pot. Excess clay from the inner wall is scraped off
and any irregularities in the clay are removed by hand or with an awl. To make
the rim the upper edge is flattened evenly, using the forefinger and the thumb,
and a thin roll of clay is carefully added and smoothed into position. To give a
very smooth finish a piece of wet calf-skin, khekhupuld, is folded over and
moved deftly round the rim. The rim profile is defined by modelling with a
thema.
The pot is covered with a cloth and taken indoors for a day or two until it is
firm enough to be inverted for the filling in of the base. If the pot is to be dec-
orated, designs are incised with an awl before the pot is taken off its support
(Fig. 5A). Red ochre, /edzogu, powdered and mixed with water, is applied to
chosen areas with a finger-tip (Fig. 5B) and burnished with a smooth river stone,
thidél6. On the following day when the pot is drier, graphite, phomd, is applied
in the same way and burnished to a metallic shine. When the pot is firm but still
pliable, it is inverted and removed from the support. The thick edges of the
open base are pinched inward and excess clay is scraped off; small pieces of clay
are added until the opening is completely closed (Fig. 6A). The base is then
smoothed with a thema (Fig. 6B) and patted into shape with a light wooden
beater, khevhad6é. At this stage unevenness may be corrected and finishing-
touches applied to the decoration. Thereafter the pot is set aside in a hut to dry
LOBEDU MATERIAL CULTURE
Fig. 4. First stages in making a pot, Modjadji’s village, 1975. A. Preparing the clay.
_ B. Positioning a rough ring of clay on the base. __C. Starting to build the wall.
D. Adding small pieces of clay.
Sy //
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 5. Decorating a pot for drinking beer, Modjadji’s village, 1975.
A. Incising the design. B. Applying ochre to the design.
for at least a week, and often for much longer, until the firing, -tshuba dibidzha,
takes place.
A number of pots are fired together except in the case of very large pots,
which are fired singly. The firing-place is a shallow depression in the ground
sited in a place sheltered from the prevailing wind. Towards sunset the pots are
positioned and supported on small stones. Slow-burning bark is stacked inside
and round the pots; kindling and old thatching-grass is piled on top and ignited.
As the grass burns, more is added until the flames are smothered and the burnt
grass is insulating the coals within. The fire is left to burn overnight. The follow-
ing morning, when the pots are cool, they are taken home. The fired ware is
usually a reddish terra cotta with some blackened patches due to reducing con-
LOBEDU MATERIAL CULTURE 59
Fig. 6. Completing the base of a pot, Modjadji’s village, 1975.
A. Filling in the opening. B. Smoothing the closed base with a
seed-pod.
ditions in parts of the fire. Burnishing may be retouched after firing (Fig. 7).
Breakages in the firing are explained in terms of poor-quality clay or the excess-
ive heat of the fire.
From the available evidence there do not seem to have been any major
changes in pottery technology over the past 40 years. The process used today
accords with Krige’s description for the 1930s (South African Museum corre-
Spondence 3.3.66) and the tools are similar to those collected by Krige in the
1930s and recorded by Lawton (1967: 172) in the 1960s. Lawton’s (1967: 176)
account of a single potter using the method of continuous coiling seems to
represent an exceptional case.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. Retouching the burnish after firing, Molototsi valley, 1978.
Types of pottery and uses
Pottery is differentiated primarily according to function which is, in general,
related to the shape, size, and decoration of the vessel.
1. Nkho (Fig. 8)
The nkho is a very large, spherical beer-pot, usually decorated on the
shoulder and round the mouth. In the 1930s these pots were used on social oc-
casions when many people were drinking beer. If a family did not have its own
nkhoé, one was borrowed from kinsmen or neighbours. By the 1970s in many
places the half 44-gallon drum had replaced, or was_used concurrently with, the
nkho for brewing beer on a large scale. In 1975 the beer-hut of the Molokwane
family contained six very large beer-pots partly embedded in the ground, two
metal drums in which the beer had been brewed, and two smaller beer-pots
from which the beer was served. By 1983 the mud-brick hut had been replaced
by a wooden shed but the large beer-pots were still in use.
2. Modzhéha (Fig. 9)
This is a large spherical pot usually decorated with bands of cross-hatching
on the shoulder and incised geometric designs round the mouth; it is used for
carrying and storing water. These pots are still much in use today as water keeps
fresh and cool in the porous earthenware. A lighter tin container is widely used
to carry water, but there is a modzhéha at every home for storing water. The
shapes of the pots from the 1930s and 1970s show great similarity, as does the
decoration.
LOBEDU MATERIAL CULTURE
Fig. 8. Beer-pot, nkhé, Modjadji’s village, 1975.
Fig. 9. Water-pot, modzhéha, Modjadji’s village, 1975.
61
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
3. Thukhwana (Fig. 10)
A fairly large, spherical or elliptical pot from which beer is served into
drinking-vessels. A thukhwana is always decorated, traditionally with incised de-
signs coloured with graphite and red ochre but nowadays also with enamel
paints. Size, shape, and design vary greatly. These and pots for drinking beer,
which are the most ornately decorated pots, are used for entertaining. In the
1930s an elliptical-shaped pot, thukhwana ya lebile (SAM-—9793), said to have
been inspired by the tread and shape of a motor-car tyre, was adopted from
Kgale potters on the highveld and became very popular among the Lobedu
(E. Krige 1977 pers. comm.). Highveld Sotho influence is also seen in the arc
design and the use of stamped lines on some of these pots (e.g. SAM-—9797,
UCT 38/83).
Fig. 10. Pots in washing-area. Beer-pot, thukhwana, left; water-pot, centre; cooking-
pot, right. Modjadji’s village, 1975.
4. Ledzhémeéla (Fig. 11A-C)
This small decorated drinking-vessel is used when offering beer individually
to an honoured guest. The smallest ones are also used to store baby food. Their
shapes vary but they are always decorated with incised, coloured designs. Many
modern examples are flat-based. ;
The ledzhéméla was introduced by Tsonga immigrants, as was the term
used to describe it (Lawton 1967: 177), and it has largely replaced the traditional
calabash beer-vessel, phafha.
LOBEDU MATERIAL CULTURE
PAE : aa
LIK DDO AS
Cc
Fig. 11. Drinking-vessels, madzhdémeéla.
A. SAM-9800, Krige Collection, 1936-8. Tradi-
tional pot for drinking beer. B. SAM-—9801, Krige
Collection, 1936-8. Flat-based pot inspired by the
shape of a jam jar. C. SAM-—10603, collected
1976. Pot of traditional shape and decoration but
with a flat base.
63
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
5. Bidzha ya khelalélé (Fig. 12)
This spherical cooking-pot is made in various sizes. The large ones are used
mainly for cooking porridge and the smaller ones for preparing vegetable relish
or meat. Decoration on these pots is minimal, usually a single line or a band of
cross-hatching on the shoulder and graphite colouring round the rim. They
blacken rapidly on the hearth and become seasoned and non-porous through
use. The simplicity of decoration and lack of red ochre distinguishes cooking-
pots from pots of similar shape not used for cooking.
a
HO we
Fig. 12. Cooking-pots waiting to be washed before use, Modjadji’s village, 1975.
6. Morifhi (Fig. 13A)
This is an open bowl used for cooking certain relishes such as termites and
locusts that do not need to be covered. It is also used as a lid for a cooking-pot
and has many other household uses. A morifhi is seldom decorated with incised
designs. Occasionally graphite is applied to the rim as a finishing-touch. |
7. Kheritswana (Fig. 13C)
|
This fairly small open bowl is used for drinking marula-cider, mokhobe, and |
for serving relish. A kheritswana used for marula-cider is always burnished with
graphite both inside and out. This distinguishes it from a morifhi of similar
Fig. 13. Bowls.
LOBEDU MATERIAL CULTURE
A. Morifhi, SAM-9706. B. Lebéda, SAM-9787
C. Kheritswana, SAM-9788. D. Lesabél6, SAM-9804. (All Krige
Collection, 1936-8.)
65
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
shape. It is often, but not always, decorated with an incised band of hatching or
cross-hatching near the rim. Modern examples are sometimes made with flat
bases.
8. Lebéda (Fig. 13B)
This large open-mouthed pot, roughly half-spherical in shape, was tradition-
ally used for steeping maize prior to stamping and during the beer-making pro-
cess. When woven beer-strainers were used, beer was always strained into a
lebéda (Krige & Krige 1943: pl. 11b) but this is no longer done, as beer is now
brewed on a much larger scale in metal drums. The /ebéda is still made and used
for soaking grain, particularly during and just after the reaping season but its use
has declined since the 1930s as many people no longer produce their own grain.
A lebéda is usually undecorated except for an incised band below the mouth.
9. Lesabélé (Fig. 13D)
This open bowl used:as a wash-basin is always decorated with a broad, in-
cised band of hatching round the rim. White chalk is rubbed into the incisions
and the bowl is burnished with graphite inside and outside. It was customary for
a wife to bring her husband a /esabél6 of warm water for washing in the morn-
ing. In the 1930s Krige recorded that these bowls were rare and had been re-
placed almost entirely by enamel basins. In the 1970s the traditional bowl was
said to be used for ritual purposes especially by mal6bo dancers.
10. Khetsikhiy6 (Fig. 14)
A deep bowl with a heavy, flat base and striated inner walls is used for
grinding tobacco, -sila leféla, into snuff (see Fig. 54). Occasionally there is an in-
cised band near the rim but most examples are undecorated.
Fig. 14. Pot for grinding snuff, khetsikhiyo,
SAM-9802, Krige Collection, 1936-8.
LOBEDU MATERIAL CULTURE 67
Snuff-making is still widely practised and involves drying tobacco leaves and
grinding them to a powder in the khetsikhiy6 with a wooden pestle, kherégélé.
The ground tobacco is passed through a wire gauze sieve or winnowed to re-
move the chaff and is then reground to a fine powder. This is blended with the
ground ash of a special plant, mokhag6 (Amarantus spinosus) to give the snuff
the desired pungency. The mixture is dampened with water to ensure a good
blend and the pot is placed on the hearth to dry the snuff.
11. Roughly made and broken pots
The Queen’s rain medicines are kept in rough earthenware pots, mehago
(Krige & Krige 1943: 273).
Roughly made pots, sometimes unfired, are used for storing seeds, dried to-
bacco or ground-nuts. These pots may represent the efforts of a novice still
learning to make pots or a poorly made pot that the maker does not want to dis-
card.
Pots cracked near the rim are often repaired with beeswax or resin
(Fig. 15). Fine wire or fibre may be bound round the rim to hold the join in pos-
ition.
Potsherds are used in a number of ways. The bowls of large broken pots are
used as basins for washing a baby (Krige & Krige 1943, pl. 5b) or when cleaning
utensils before use. Smaller sherds are useful as containers for mixing ochre,
graphite, or herbs. New pots are built up on flat potsherds. Certain medicines,
especially those burnt to give ashes, are kept on sherds. Small pieces of broken
pottery are embedded in the floor of the courtyard to act as door-mats and to
prevent erosion in drainage areas. Sherds were also embedded in the bases of
threshing-pits in the fields.
Fig. 15. Pot repaired with resin, Modjadji’s village, 1975.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
12. Pots used in termite traps
In the early summer months traps, involving the use of small pots, are made
to catch dinntwa termites (a very important food source). A hole, mokhobd, is
dug in a termite heap, kheulu, and a pot placed inside. A framework of sticks,
madzwari, is placed over the hole and covered with leafy twigs. A small opening
is left to allow light in near the pot. When the termites swarm they fly toward
the light and get trapped in the pot.
Other uses of clay
Toys
Clay is used by children to model toys. Herd-boys make clay oxen to resem-
ble the long-horned cattle they admire. Small girls make miniature clay imita-
tions of their mothers’ utensils and use them in play-housekeeping which is an
informal training in home-making.
Tuyeres
When iron-working was an active industry, clay was used to form the
nozzles for the bellows and for constructing the walls of the furnace.
Whorl
A ceramic whorl was used to weight the spindle used in spinning wild cotton
(see p. 91):
GRASSES AND OTHER PLANT FIBRES
Formerly the indigenous vegetation provided a very rich source of grasses,
sedges, creepers, barks, and other plant fibres. By the 1980s much of the natural
vegetation had been cleared for cultivation with the result that the plant re-
sources had been reduced and greater distances had to be travelled to find par-
ticular plants. Krige & Krige (1943: 47) commented on the impressive body of
botanical knowledge that the Lobedu put into practice in their craftwork. Crafts-
men had specialized, practical knowledge of the many different plant fibres suit-
able for making string, baskets, and mats. The quality of the material was
carefully selected for the object in mind. Pliability was necessary for strainers,
strength for baskets, and resilience for sleeping-mats. The long-established use-
fulness of certain plants is reflected in the vernacular terminology. The species
of acacia most frequently used for weaving winnowing-baskets is known as mo-
loha, from -loha meaning to weave. Many objects have the same name as the
material used to make them, for example the sleeping-mat, pate, made from the
sedge of the same name. Symbolic associations also influenced the choice of ma-
terials especially in the case of initiation costumes.
Seasonal availability of grasses and sedges restricts their use to certain times
of year. Thatching-grasses may not be cut before Modjadji has given her con-
sent. This restriction reflects the chief’s control over an essential resource but it
LOBEDU MATERIAL CULTURE 69
is also a practical measure ensuring that the grass is not cut until ripe (Krige &
Krige 1943: 47). Ease of collection influences the choice of material. If the ap-
propriate material is inaccessible because of drought or distance from the
source, a suitable substitute may be used. Substitute materials, however, do not
usually have the particular qualities of the original material and this is evident in
the product. The accessibility of cultivated sisal as well as the strength of the
sisal fibre has made it an acceptable substitute for many of the indigenous fibres
formerly used for the making of rope and string.
Basketry techniques
The two main technical categories of basketry, coiled or sewn and woven,
and the variations found in Lobedu basketry are defined briefly below.
In coiled and sewn basketry a coiled foundation is sewn together with one of
a variety of stitches. A needle may be used but is not necessary if the sewing-
fibre is fairly rigid and is used in conjunction with an awl. In simple oversewing
each stitch passes over the foundation coil and pierces the row below. If the
stitch splits the stitch in the previous row a forked effect is produced, this is
known as furcate sewing. Another variation, called bee-skep, has widely spaced
stitching connecting the coils at intervals, so that each stitch is just above the
one below. The foundation material is variable and may consist of a single rod
known as a simple foundation or a cluster of strands known as a multiple founda-
tion. If the material of the coil is too stiff to allow a stitch to pass through it, the
coil is joined by passing a strand through the fabric of the basket to link adjacent
rows of the coil. A sewn technique is used for certain mats and beer-strainers in
which case the work is flat or cylindrical and the sewing-strand passes through
the other element.
In woven basketwork two elements, the warp and weft, cross over and
under each other. The manner of crossing or weaving creates different patterns.
A twill, which is widely used for making winnowing-baskets, is characterized by
the weft passing over and under two or more warps with each successive row
starting one warp ahead of the previous row. Other woven techniques used by
the Lobedu include wicker-work in which the warps are rigid and the wefts flex-
ible (as in a reed-door), and twining in which two wefts pass alternately in front
of and behind each warp (as in sleeping-mats).
In practice both woven and sewn techniques may be used in a single basket,
for example the rim of a woven winnowing-basket is oversewn. When used with-
out qualification, the terms apply to the main fabric of the basket.
Uses of basketry
Baskets
1. Coiled, sewn baskets
The mothagha or kherodo (Fig. 16A) is a large basket for carrying produce
from the fields and for general household use. A coiled technique is used in its
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 16. Sewn baskets. A. Mothatha, SAM-9738. B. Khethathana, SAM-9741. (Both
Krige Collection, 1936-8.)
manufacture, and the material most often used for both the coil and the sewing-
strand is the stem of the creeper lebibye (Cocculus hirsutus). To prepare it for
use, the creeper is boiled in water until the outer bark is loose and can be pulled
off easily. While still damp and flexible from soaking in warm water, the stem is
cut through the middle and stripped of the inner pith (Fig. 17). The stems can
then be scraped down to the required thickness for use. For the foundation it
may be left whole or split in half; the wrapping-fibre is pared down into thinner
strands. Lebibye may be prepared well in advance and stored until required.
The mothatha and similar smaller khethathana baskets (Fig. 16B) are made
LOBEDU MATERIAL CULTURE 71
Fig. 17. Women preparing strips of /ebibye. Note pot of water for soften-
ing the fibre and the use of the toes for holding the fibre. Photo: E. J.
Krige, 1936-8.
by women (Fig. 18). The foundation is joined by passing the thinned creeper
over two coils to interlock with the previous row (Fig. 19A). An awl, morogola
(see Fig. 110B), is used to make a space for the creeper to pass through. Dry
lebibye is inflexible and must be dampened with warm water before use, as is
generally the case in much basket-work. The rim of the basket is thickened with
any pliable stick and oversewn in a herring-bone pattern (Fig. 19B—C). A piece
of hide may be sewn onto the base to protect it from wear. The final product is a
very strong durable basket.
a2 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 18. Woman working on a mothatha. She is using an awl to make a
hole for the sewing-strand to pass through. Photo: E. J. Krige, 1936-8.
The smaller domestic basket, khethathana, is generally made in the same
way but not always of the same materials. Bark fibre may be used for the sew-
ing-strand but this is not as durable as /ebibye as it frays when cleaned in the
customary way of scouring with sandstone. If grass is used for the foundation, as
in one example (Fig. 16B), the coil is many stranded and is sewn together, in
this case with bark fibre, using a furcate (Fig. 20) stitch. This basket lacks the
firmness and strength of those made with /ebibye.
LOBEDU MATERIAL CULTURE
Y. BRANCO
Cc
Fig. 19. A. Interlock oversewing-technique. B. Herring-bone
binding. C. Rim attachment to basket.
= 2V2\ oe [NINN
Be
=v: VEO VR
VY. BRANCO
Fig. 20. Furcate sewing-technique.
He:
74 ANNALS OF THE SOUTH AFRICAN MUSEUM
Formerly, when there was a surplus of grain to be stored, men made large,
coiled grain baskets with lids that could be sealed with dung. A kheséh6é was
made with a foundation of lefhe grass sewn together with bark fibre using a bee-
skep stitch (Fig. 21). The rim of the mouth was oversewn with bark fibre in her-
ring-bone stitch.
Fig. 21. Beeskep sewing-technique.
2. Woven baskets
The woven winnowing-fan, lesél6 (Fig. 22A), the smaller but similarly
formed basket, tsana, and the serving-basket, kherodwana (Fig. 22B) were
made, and in the case of the Jesél6 and tsana still are made, by men from thin
strips of wood cut from a branch of a suitable tree, most commonly moloha
(Acacia ataxacantha). Their manufacture is described here and not in a later sec-
tion on the use of wood as, apart from the cutting of the wood, basketry tech-
niques are employed.
In addition to the wood slivers used for both warp and weft elements,
lengths of firm but flexible wood and creeper stem are required for the rim. The
materials are collected and prepared in advance. The strips of wood are pared
with a knife to similar lengths and thickness; the wood for the rim is cut to the
length that will allow it to be bent into a hoop, legdk6, the size of the circumfer-
ence of the rim, and the overlapping ends are secured with a small dowel. The
creeper lebibye is prepared as described above for the coiled baskets.
The craftsman first dampens the wood slivers in warm water, takes four
strips to start and incorporates additional strips by weaving in a straight twill
over two (Fig. 23A) until a square is complete. While weaving the craftsman
supports the work on his lap or on a piece of wood on the ground. The hoop is
tied to the flat woven square in four places, the corners of the square are cut off
and the remaining circular piece is then pushed through so that the hoop is on
the outside. A thin plant stem, khuzwane, is tied on the inside of the rim, which
|
'
|
|
.
/
LOBEDU MATERIAL CULTURE 75
Fig. 22. Woven baskets. A. Winnowing-basket, Jlesélé,
SAM-9679. B. Serving-basket, kherodwana, SAM-9673.
(Both Krige Collection, 1936-8.)
is then oversewn with the creeper binding (Fig. 23B—C). An awl, morogola, is
used to make holes through which the creeper is threaded (Fig. 24A). The ends
are worked into the binding and the finished product is very strong and neat.
Winnowing-baskets collected in the 1970s do not differ from those collected
by Krige in the 1930s and they are used whenever it is necessary to separate
coarse from finer matter. Winnowing, -fhefhéra, is essentially an up-and-down
fanning movement during which lighter matter is separated from heavier. It is a
— ee
——_—_——7;
—_— =
TU
LOBEDU MATERIAL CULTURE
Fig. 24. A. Basket-worker binding the rim of a /esél6. Note use of awl. Mohale’s village, 1973.
B. Relish drying on an old /esél6, Modjadji’s village, 1975.
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
process much used by Lobedu women both in the fields to free the chaff from
the heavier grain (Figs 25-27) and at home when stamping maize into meal or
when they wish to separate dry ingredients.
When maize is stamped by hand the J/esél6 is a necessary complement to the
pestle and mortar in the successive stages of the process that produces meal of
different texture and so adds variety to the staple porridge diet. After the first
stamping, -thovola, the grain is winnowed in an up-down action, -fhefhéra, to
separate the outer husks, which are made to fall to the ground. Thereafter the
basket is moved in a rotating action, -séléla, so that the larger husks come to the
Fig. 25. Woman winnowing sorghum with an up-down movement,
-fhefhéra, to separate the grain from the husks. Photo: E. J. Krige,
1936-8.
LOBEDU MATERIAL CULTURE 79
Fig. 26. Woman holding the winnowing-fan to the wind to separate the
heavier grain from the chaff. Photo: E. J. Krige, 1936-8.
top in the centre. These are removed by hand and kept as food for the pigs; the
de-husked grain is returned to the stamping-block and the process is repeated at
least twice. The husks of the next winnowing are kept for making mabudu, a
light beer; those of the following are used for making maase, a fairly granular
porridge. At this stage the stamped grain, thovole, is placed in a pot, covered
with water and left for a number of hours before being drained on a /esél6. The
water in which it was steeped is kept for making soft porridge, vhothithi, which
has a slightly sour flavour. The grain is then ready for the second stamping-
process, -seda, which results in the fine, white flour used for making the staple
porridge, vhoswa.
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 27. Woman separating the coarser grain in the centre of the basket, after winnowing with a
rotating action. Photo: E. J. Krige, 1936-8.
Apart from its use in winnowing, the Jesé/6 can be used as a tray for serving
guests at beer-drinks and for many other general household purposes, including
the drying of relish (Fig. 24B) for use in the dry winter months when fresh
greens are scarce. The smaller tsana has many domestic uses and is especially
useful for holding maize-meal or relish during the preparation of food, and as a
lid for other utensils.
The kherodwana is a waisted basket with a lid, that was used traditionally
for keeping food warm, for serving an honoured person, and also for carrying
goods. The lid, made in the same way as a tsana, could be used on its own as a
small basket. The shape of the basket was designed to fit a wooden dish, delod,
on which porridge was served and set aside in the basket to keep warm until
eaten. The base of the kherodwana was made in the same way as a lesélé and
similar materials, strips of moloha wood and ditsidwa or lebibye creeper, were
used. For the sides of the baskets the ends of a rectangular mat of twilled wood
slivers were woven together to form a cylindrical shape that narrowed at the
lower edge to fit into the rim of the base. The join was oversewn with creeper
binding. The upper edge of the basket was fitted with a ring of wood as for the
rim of a lesél6 and was oversewn. The making of this basket required more
technical skill than other baskets and it had particular social importance.
LOBEDU MATERIAL CULTURE 81
Fig. 28. A. Woman making a twined sleeping-mat. Photo: E. J. Krige, 1936-8. B. Woman
making a twined sleeping-mat. Ramalepe’s village, 1973.
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the 1930s every Lobedu bride took a kherodwana to her new home as a
symbol of honour to her husband. It was customary for the bridegroom’s father
to instruct the bride, through his wife, to serve food to the groom in this ‘mar-
riage’ basket (Krige notes). By the 1970s changing social conditions had made
the kherodwana obsolete. This followed changes in the pattern of marriage re-
lated to an increase in migrancy. As it became common for young men to be
away for long periods of migrant labour, many marriages were contracted in the
absence of the bridegroom. In these circumstances there was no need for the
customary use of the kherodwana in marriage ceremonial or in the domestic
sphere.
Mats
Two types of mat were used: pafe, a twined sleeping-mat, and khegdyo, a
_sewn mat for drying meal or for sitting on. Both mats were formerly made only
by men (Krige notes) but as migrancy reduced the number of men in the villages
women took over the task of making sleeping-mats. In the 1930s both men and
women wove mats (Figs 28A, 29), in the 1970s they were made almost exclu-
sively by women (Fig. 28B).
Fig. 29. Man making a twined sleeping-mat. Note use of pegs and pot of water for softening
the fibres. Photo: E. J. Krige, 1936-8.
LOBEDU MATERIAL CULTURE 83
LA EZ
V. BRANCO
Fig. 30. Techniques of finishing edges of mats. A. Whipped edge of sleeping-mat. B. Knotted
edge of twined sleeping-mat. C. Wrapped edge of sewn grain mat.
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
Sleeping-mats are made from a sedge, pate (a species of Cyperaceae). Four
or five lengths of sedge are tied together to form multiple warps, and work pro-
ceeds from the one side or the centre by twining the warps together with string,
the ends of which may be secured with pegs (Fig. 29). The string is usually made
from sisal, mokhdéda, but formerly molete (Triumfetta pilosa) bark was used.
Rows of twining are evenly spaced across the width of the mat. The ends of the
sedge are normally trimmed and finished off with a simple knot (Fig. 30B). The
ends of the string are wound round the last warp a few times and knotted. Warp
ends may be whipped with single-ply twisted sedge (Fig. 30A). The completed
product is a thick and comfortable mat on which to sleep.
Making sleeping-mats on a vertical wooden frame (Fig. 31) is a recent prac-
tice which seems to have been introduced by Tsonga-speakers. Grooves are
made at regular spaces along the horizontal beam of the frame to mark the posi-
tion of the strings that are tied to the first length of sedge. Work proceeds by
_ adding bundles of sedge that are twined together with the strings until a mat of
the required length is made. During the work the strings are weighted with
stones to keep the mat in position and the tension even. This is a much quicker
way of making mats than the traditional method.
The khegdyo mat for drying meal after stamping is no longer in regular use
as most people now buy their maize-meal. It was made from lengths of thin
sedge, khegdyo (Cyperus textilis), sewn together by a strand of twisted molete
bark fibre passed through the sedge with a metal needle, /emao, in a series of
rows. The edges were trimmed and wrapped with string (Fig. 30C). According
to Krige (notes) this type of mat was adopted from Tsonga immigrants.
Fig. 31. New method of making a sleeping-mat on a wooden frame. The twining-threads are
weighted with stones. Molototsi valley, 1973.
LOBEDU MATERIAL CULTURE 85
Wicker doors
A door, khezwala vhatimana, made by men using a wicker basketry tech-
nique, was the traditional door for a cooking-hut. Translated literally, the ver-
nacular term means ‘something by which selfish people keep each other out’, i.e.
to obviate the need for hospitality.
These doors were constructed by making a lattice of thick mbaseriti reeds
and any suitable thinner poles or sticks held together by wrapping (Fig. 32) with
a strong bark fibre such as motswiriri (Bauhinia galpinit).
CCM
D
Cm
bil
( i
aa T
Ye
i
——
y.BRANCO
Fig. 32. Wicker technique formerly used for making doors.
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
Reed doors were not hinged. To close one from the inside a pole was
passed through a thong loop attached to the centre of the door, and across the
door frame; from outside it was held up with a heavy object such as a wooden
pestle. These doors were smeared with cow-dung both to prevent draughts and
to protect them from infestation by weevils. When not in use as a door it could
be used as a large rigid mat on which to place things. Krige records that they
were rare in the 1930s; they were not seen at all in the 1970s.
Strainers
The traditional Lobedu beer-strainer lethéd6 (Fig. 33A) was woven by men
(Fig. 34) from a number of different plant fibres, for example ‘bobbejaanstert’,
Fig. 33. Beer-strainers. A. Woven, SAM-9750. B. Sewn, SAM-9749. (Both Krige Collection,
1936-8.)
kheruje (Vellozia retineris), tsenyane grass (Ophioglossum sp.), or strips of wood
from the molete or moloha-thédo tree (Triumfetta sp.). The same material was
used for both warps and wefts, which were woven together in a twill pattern.
The start was made by knotting a row of strands together (Fig. 35A) and weav-
ing continued in a tubular manner. To close the base the strands were gathered
tightly, the ends knotted and turned inside the strainer.
A sewn strainer lethéd6 la horokiwe (Fig. 33B), said in the 1930s to have
been recently adopted from Tsonga-speakers (Krige notes), was made from the
midrib of the leaf of modolo (Ensete ventricosum) twisted into two-ply string.
The start was made at the open end and the sewing-strand was passed through
LOBEDU MATERIAL CULTURE 87
Fig. 34. Man weaving a strainer while attending a court case in the
khoro. Photo: E. J. Krige, 1936-8.
knots in the string (Fig. 35B) to form a circle. A needle was used for the sewing
and work continued in a spiral with the sewing-strand passing through the string
(Fig. 35C). At the base the strands were sewn into a tight bunch.
Both woven and sewn strainers were used in the same manner. Unstrained
beer was poured through the /ethéd6, which was wrung in the hands with a
twisting action so that the liquid passed through the fibres and the dregs re-
mained behind. The receptacle for the strained beer was a wide-mouthed pot,
lebéda (see p. 66). By the 1970s both types of basketry strainer had been re-
placed by a large wooden sieve, sef6 (see p. 113). This replacement related to an
increase in the scale of brewing beer for sale.
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LOBEDU MATERIAL CULTURE 89
Playthings
In summer when the grasses are long, boys and girls make a variety of play-
things and ornaments, using basketry techniques. Examples in the Krige Collec-
tion include armbands, a grass peaked cap, képisi ya bosigijane, made in
imitation of a schoolboy’s cap, and a hat, gadiba, also made of bosigijane grass
(Sporobolus pyramidalis). The hats have coiled foundations of bundles of grass
sewn together with single strands of grass. Plaited and woven techniques
(Fig. 36) are used for making armbands that imitate the ornaments worn by
Fig. 36. Plaiting- and weaving-techniques used for
making armbands.
adult women and are referred to by the same names, maséga and khefhddo.
Single lengths of grass are made into toy arrows to be used with a small bow
made of a pliant stick and bark.
Other uses of grass and plant fibres
Brooms
The simplest broom, J/efiélé6 (Fig. 37A), is easily made from a bundle of
lefiél6 grass (Aristida congesta) pulled out by the roots and tied at the grip with
string. Rough brooms for sweeping outside and for cleaning the hearth are made
in the same way from motataile (Arthrixia phylicoides) or lefaladza maru (As-
paragus phimosus). When bosigijane grass is used the broom is made in a differ-
ent way (Fig. 37B) as this grass does not pull out easily by the roots. Lengths of
grass are twined together, folded over, rolled up, and bound at the grip with
90 ANNALS OF THE SOUTH AFRICAN MUSEUM
Manin
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cee
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Fig. 37. Two types of broom. A. Lefiél6 made of grass pulled out by the roots. B. Grass
twined and bent over before being bound to form grip of broom.
string. Grass brooms are still made by women during the season when the grass
is available, and they are in general household use.
Cordage and spun thread
Lenti is string or cord made from plant fibres. It is made by both men and
women but traditionally it was made mainly by men.
The choice of specific barks, creepers, fibrous leaves or roots depends on
the intended use. The quality of the fibre is taken into account and it is prepared
accordingly. Certain bark and bast fibres, for example moga (Acacia rehman-
niana), produce very strong lashings suitable for structural work; other fibres,
such as molete (Triumfetta pilosa) have pliability suitable for basketry. Strips of
palm or banana leaf make adequate temporary lashings for tying bundles but
would not be durable enough for use in hut-building. Other fibres recorded for
making ropes and string include mokhéba (Dombeya rotundifolia), mothanari
(Colophospermum mopane), motswiriri (Bauhinia galpinii), mofwara tsweni
LOBEDU MATERIAL CULTURE 91
(Grewia sp.), mothala malagani (Acacia karoo), and mokhéda (Sansevieria sp.).
Many barks are cut into strips and used as lashings without further prepara-
tion. If collected in advance, the strips are rolled into balls or bundles and stored
until required. Before use, dry brittle bark-fibre is soaked in water or may be
boiled to strengthen it.
String is made by separating the fibres of the chosen bark or plant and twist-
ing the fibres together by rolling them by hand on the thigh or calf (Fig. 38). The
Fig. 38. Woman rolling sisal string, Modjadji’s village, 1975.
deftly executed rolling action, -hofa, is in two directions, both backward and for-
ward. Two or three lengths of string may be twisted into two- or three-ply cord,
following the same method.
The introduction of cultivated sisal, Agave americana (also mokhéda),
made it more readily available than the indigenous fibres and this, together with
the ease of working with the fibres, has resulted in its replacing most other ma-
terials for making rope and string.
Formerly wild cotton, /egudu (Gossypium herbaceum var. africanum)
(Fig. 39A), was spun by men into thread and plaited into girdles or necklets by
women. Krige records that in the 1930s cotton was spun on spindles
(Fig. 39B—C) weighted with ceramic whorls, kheritswana, and that there was
considerable demand for handspun thread, Jedede or letshida, which was used
for the ties on skin skirts and baby-slings as well as for necklets. By the 1970s
bought thread had replaced handspun cotton, spinning was remembered only by
a few old men, and pieces of handspun thread were treasured as heirlooms.
92 ANNALS OF THE SOUTH AFRICAN MUSEUM
B
Fig. 39. A. Wild cotton growing near Modjadji’s
Nek, 1973. B. Lobedu man spinning cotton.
Photo: E. J. Krige, 1936-8. C. Spindle with
ceramic whorl, SAM-9772, Krige Collection,
1936-8.
LOBEDU MATERIAL CULTURE 93
Headpads
Bundles of grass shaped into the form of a ring and bound roughly with any
plant fibre such as grass, bark, or banana leaf are used to support heavy loads
carried on the head. A headpad, khare, may also be made of a rolled-up piece
of cloth serving the same purpose.
Nests for hens
A nest, khetutu, for hens is made by coiling and sewing bundles of moha-
lana grass (Rhynchelytrum repens) to form a sphere and cutting out an opening
for the entrance. Nests would be placed on a high platform to keep the hens out
of reach of predators at night. The keeping of hens is a Tsonga practice that has
been gradually adopted by the Lobedu but not without some resistance. This
was strongest in the royal village and even today, although many people else-
where in the Lobedu area keep hens, they may not be kept at the capital.
Another kind of hen-house, recorded in the 1930s, was a small, conical
grass structure bound with grass at the apex. A stone was placed over the open-
ing to close the entrance. This was not seen at all in the 1970s.
Thatching
Thatching is a seasonal activity that takes place after June when the grasses
are ready to be picked and before the rains start in late October. Women gather
the grasses in large bundles (Fig. 40A) and men do the thatching. The old way
of doing this was simply to lay the grass on the roof framework of closely packed
rafters, to tie down the first layer of thatch at the lower edge so that the roots
pointed downwards, and thereafter to place the other layers of grass on top with
the roots facing in the opposite direction so that they interlocked with the lowest
layer. Letshumelo grass was used for this kind of thatching (Krige 1982 pers.
comm.). Withies or grass rope may be placed on top to hold down the thatch
more firmly (Fig. 40B).
The modern thatching-technique is to sew the grass on to the rafters with a
long needle, /emao, made of a hard wood such as moréjé (Dicrostachys cinerea).
Work starts at the apex of the roof. Thazi grass (Hyparrhenia filipendula) is con-
sidered the best grass for the first layer as it has a smooth stem that does not
break easily. Thereafter other grasses are placed on top and sewn down. By sus-
pending a small ladder from the roof pinnacle the thatcher can move around on
the roof quite easily (Fig. 41). The sewn thatching-technique is said to have been
adopted from the method used by white farmers.
Vuhwera and vyali costumes
Young men in the vuhwera (North Sotho: bogwéra) initiation, which is held
concurrently with the girls’ vyali (North Sotho: baale) (for full description see
Krige & Krige 1943: 126-140), are taught by older initiated men how to make
94 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 40. A. Woman carrying thatching-grass, Modjadji’s village, 1973. B. Old type of thatched
roof, Modjadji’s village, 1975.
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96 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 42. Mohwera dancing in courtyard. Photo: E. J. Krige, 1938.
indigenous fibres, there is an awareness among conservative Lobedu of the sym-
bolic importance of the traditional materials.
The following information was given by Simeon Modjadji in 1975. There
are two types of vuhwera costume—a light one, mamegwane, worn when out in
the bush hunting, and a heavier dancing-costume, thégwa. Both have loose
fringed skirts, shoulder-bands and a head-dress but the dancing-costume is far
more elaborate and has row upon row of grass cross-bands forming an immense
chest and shoulder covering on which rests a crested head-dress adorned with
feathers or fur (Fig. 43). Armbands, anklets and, in some cases, underskirts are
made of lengths of grass twined tozether to form flat bands. Thazi grass (Hypar-
rhenia filipendula) which has a smooth glossy stem, is the basic material used for
all parts of the costume except the skirt, which is made from strips of molala
palm leaf. Two-ply fibre string is used to join the pieces of grass together in a
special twining technique referred to as khavhe (Fig. 44B). This kind of twining
is used for the shoulder-bands, mofhaga, the arm- and legbands, zwifhddo, the
head-dress, thohd, and for all other parts of the costume except the skirt, mod-
javhe. This is made by knotting long strips of palm leaf on to a rope girdle that,
when worn, is wrapped round the waist in layers.
Towards the end of the vuhwera the initiates spend most of their time going
about to dance at different villages on the invitation of the headmen, or at the
Se fe eS ea
LOBEDU MATERIAL CULTURE 97
Fig. 43. Elaborate costumes worn by the magdégébya during the 1938 vyali-vuhwera initiation.
Photo> E~J> Krige, 1938:
capital to pay respect to Modjadji. The costumed dancers perform in pairs or
singly, accompanied by drumming and singing of the vyali girls. During the vu-
hwera dancing the skirt flaps rhythmically, but the weight of the entire costume
restricts the dance to slow graceful movements. The wearing of the heavy cos-
tume abrasive to the skin of the wearer is part of the ordeal that initiates must
endure. The overall effect is an imposing spectacle much enjoyed by the on-
lookers who reward their favourite initiates with gifts of ornaments or snuff.
In the 1938 initiation the most spectacular costumes were worn by the vah-
wera of Rabothada whose performance at the capital traditionally closed the ini-
tation. Their dancers, the magdgébya (Fig. 43), had magnificent costumes with
head-dresses surmounted by animal figures, and had underskirts trailing the
ground with dramatic effect. According to Krige (1982: 8) these costumes rep-
resented the finest weaving-skills.
The costumes worn in the vuhwera of 1974, which was the next school held
on a national scale after 1938, displayed a deterioration of weaving-skills. The
elaborately costumed magégdbya did not appear at all as it was said that no one
knew how to make the costumes (Krige 1982: 8). The vuhwera costumes worn in
schools run independently by headmen, but with the permission of Modjadji,
also revealed a decline in craftsmanship.
ANNALS OF THE SOUTH AFRICAN MUSEUM
B
Fig. 44. A. Mohwera dancing to accompaniment of vyali girls, Molototsi valley, 1973.
B. Detail of technique used in mohwera costume.
LOBEDU MATERIAL CULTURE 99
Vyali girls do not wear elaborate costumes at any time during the initiation.
During the early stages they wear only short wraps round the hips. Formerly
these were made of hard, unworked skins but in the 1938 initiation cotton cloth
was substituted (Krige & Krige 1943: 133) and has continued to be used instead
of skin. At a later stage the girls wear bandoliers (Fig. 44A) plaited from Jetate
grass and, according to Krige & Krige (1943: 136), dancing skirts, dali, were
made of a special kind of reed strung together in pieces about 2,5 cm long; they
were used at both the 1938 and 1974 vyali (Krige 1982: 9).
An essential aspect of the vyali and vuhwera is the revealing of digdma,
mysterious figures or objects unknown before initiation. These have not been
seen by the writer but, according to Krige & Krige (1943: 135), in 1938 many of
the digOma were masked figures in costumes made of grass, reeds or leaves.
These masked figures were connected with mummery or verbal formulae, each
with an esoteric meaning.
Khiudogane, the Bird of Muhale, is the greatest géma and ruling spirit of
the vyali. It appears in the moonlight wearing a strange costume consisting of a
conical wooden framework similar to the structure of a hut roof. From this
framework, which rests on the shoulders of the crouching wearer, hang long
strings of blackened fibre. The fibre of the moga tree (Acacia rehmanniana),
which becomes black when soaked in water, was traditionally used but more re-
cently this has been replaced by sisal fibre blackened by rubbing with charcoal.
The knot by which the strings are attached to the framework is the same as that
used to attach the fringes of the initiation skirt to the girdle (Fig. 45). At the
pinnacle, thoho, there is a small sphere of wood coated with black resin in which
red seeds and a feather are embedded. The Bird is believed to come from a pool
in the river and the vyali girls sing its praises even when it does not appear in the
moonlit courtyard. It may communicate its commands at a distance by whistling
through a special instrument, mphenyi, made of a reed pipe stopped at one end
with a membrane from a spider’s web (Krige 1982: 27). The Bird arrives only
when food is plentiful and may be seen as the embodiment of fertility (Krige &
Fig. 45. Technique used for attaching skirt fringe to mohwera costume.
100 ANNALS OF THE SOUTH AFRICAN MUSEUM
Krige 1943: 135, 139-140). The entire vyali, which spans a full agricultural
cycle, is at one level an initiation for girls and at another level a fertility and rain
rite.
For comparative data on the Kgaga bogwéra and baale initiation schools,
which have many elements in common with the Lobedu, see the recent analysis
by Hammond-Tooke (1981: 55-82).
WOOD AND REED
The indigenous environment is rich in trees and traditionally provided a
wide range of woods for carving, for constructing huts, granaries and livestock
enclosures, as well as for fire-wood.
Integrated with a practical knowledge of the natural vegetation, was the Lo-
bedu concept of order in nature as something controlled through the power of
medicine and magic, through the ancestors, through the divinity of the Queen,
and through the mysteries associated with fertility (Krige & Krige 1954: 68).
Certain woods used mainly for medical or magical purposes, for example woods
used for the protective wand buried across the entrance to a village or for the
medicated pegs around the perimeter, could not be brought into the village for
secular use as this would reduce their supernatural potency. The same taboo ap-
plied and still applies today to the trees associated with shrines and the wood
from which charms, dithugula, are made (Krige & Krige 1943: 167). The movil6
(wild medlar, Vangueria infausta), the moludu (white stinkwood, Celtis africana)
and kherale (Gardenia jovistonantis) are widely believed, not only among the
Lobedu, to have magical properties (Palmer & Pitman 1972: 203). Kherale was
one of the woods used for the medicated pegs (National Open Air and Cultural
History Museum, ET 62/7, collected near Duiwelskloof) which were used, to-
gether with medicines, to protect villages from evil spirits. Twigs from the khadi
creeper (Adenia gumnifera), which contains much moisture in its stems, were
used for making brushes for sprinkling ‘cooling’ medicines in rituals of purifica-
tion (Krige & Krige 1943: 274).
Symbolic associations of certain trees are important in understanding their
uses. The vitality, luxuriant growth and abundant fruits of the Celtis africana are
positive qualities that give the tree protective powers. The lush fruits of the mar-
ula (Sclerocarya caffra) and the wild-fig (Ficus sansibarica) are the epitome of
fertility, apart from being edible and rich in essential proteins. An infusion of
Cussonia spicata, a species with swollen underground roots, is used by Lobedu
mothers to make their babies fat and strong. The great wild-fig trees shading
shrines have ‘cooling’ properties, as do the succulent bulbs used in the rite for
removing the impurity or ‘dirt’ (khetshila) of death (Krige & Krige 1954: 69).
The Daja forest in which royal chiefs are buried is held sacred and it is a
serious offence to cut wood there (Krige 1931: 210). This taboo applies also to
Moholwe where the sacred drums are kept and, in general, to all burial sites.
The felling of trees here would cause the anger of the ancestors. Certain mem-
bers of the community who have died unnaturally are buried in wet soil under
LOBEDU MATERIAL CULTURE 101
overhanging trees near rivers, in order to ‘cool’ their ‘heat’, which contaminates
the earth and causes drought. If the trees around these graves are desecrated,
even by accident, a ritual of purification must be performed. These beliefs are
still held even among converts to Christianity as shown by a purification ritual,
phuphutshéla madiba, witnessed in 1973. A schoolteacher, who had accidentally
started a fire that spread to a burial site and burnt many trees, had to sacrifice a
sheep and be sprinkled ritually with ‘cooling’ medicines containing, among other
things, rain-water and the undigested stomach contents of the sheep. A bunch of
khadi twigs was used as a brush. The burnt graves were also treated with medi-
cines. This was necessary to appease the ancestor spirits and prevent them from
sending wind that would dispel the rain clouds.
Although important in understanding the Lobedu concept of their natural
environment, taboos on the use of specific woods did not seriously restrict the
technology as they applied only to a small fraction of the many available woods.
Wood-working techniques
‘For each of the crafts—hut-building, fencing, making stamping-blocks,
sledges, hoe-handles, milk-pails, spoons, platters, winnowing-baskets, drums
.. . different qualities in the wood are regarded as important’ (Krige & Krige
1943: 47). Morididi (leadwood, Combretum imberbe), for example, was known
to yield charcoal of great enough heat for smelting iron, and mbidibidi (Ekeber-
gia capensis) was valued for its soft light wood that was easy to carve. Hard, ter-
mite-resistant woods such as kiaat and mopane were selected for structural work
and, although difficult to carve, for pestles and spoons that would wear down
too quickly if made of softer wood. Even the everyday task of collecting fire-
wood for the hearth required a knowledge of which woods burnt quickly or
slowly, which smoked a lot, and which would produce long-lasting coals.
The choosing of a suitable wood was, in fact, one of the most skilled aspects
of wood-working. The preparation of wood for carving and the actual carving
techniques were, by comparison, less demanding. Wood was seldom seasoned be-
fore use and most objects were carved from the solid without need for joinery.
The traditional tool-kit consisted of an adze, bédwana, a poker, mor6é, and
a number of gouges, mah6r6, of varying size each handled in a particular way
(Fig. 46). Before contact with white traders, iron was obtained through trade
with neighbouring people, notably the Phalaborwa and the Venda, and tools
were forged locally. For many years now scrap metal, motor-car springs and files
have been heated and hammered into blades for tools and hafted in the old way.
More recently modern tools have come into use and axes, saws, chisels, pen-
knives, and drills are used by those who can afford to buy them. The traditional
tools, however, were well designed for their many uses and they have not been
replaced. It is common to find hand-made tools used concurrently with bought
tools (Fig. 47). Modern tools have made the felling of large trees much easier
than in the past when it was necessary to make a fire at the base of the trunk and
gradually cut through the burnt layers of wood.
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 46. Woodcarver making a dish, de/6, using a long-handled gouge
with a bent blade. Note the characteristic position in which he is sitting
while supporting the dish against a log of wood and holding it with his
feet. Photo: E. J. Krige, 1936-8.
Traditionally there was some degree of specialization in the making of ob-
jects that required particular skill, but in general most men could make domestic
utensils and handles for tools (Krige & Krige 1943: 50). Even skilled craftsmen
were not full-time specialists and they seldom specialized in making only one
type of object. It was not uncommon, however, for a man or a family to win a
reputation for excelling in making particular objects.
Drums and mortars that required the felling of a tree were usually only
made to order by craftsmen who possessed the necessary tools and expertise.
LOBEDU MATERIAL CULTURE 103
Fig. 47. Traditional and modern tools used by Calvin Ramatswalela, 1973. Left to right:
purchased axe, traditional adze and three gouges, modern chisels, files, and knives.
Smaller objects such as dishes and spoons were more generally made. If a man
had time to carve in excess of his own needs or was commissioned, he would ex-
change wooden objects for grain or sell them at the local market. Seasonal activ-
ity regulated both the practice of the craft and the demand for specific products
(Krige & Krige 1943: 32). At the peak of the agricultural season craft-work
came to a standstill and during the latter part of the dry season rebuilding and
rethatching of huts took priority over other craft activity.
Most wooden utensils had a relatively long life-span and seldom required
replacement unless attacked by termites. In the absence of a developed trading
system there was no stimulus for surplus production and local demand was not
great enough to support full-time specialists.
Since the 1930s, wood-working has become less generally practised. Restric-
tions on the felling of most indigenous trees has limited the legal use of certain
woods and the pole-and-daga method of hut-building is no longer used. The
demand for wooden stamping-utensils declined as the economy became increas-
ingly dependent on purchased maize-meal. Furthermore, migrant labour re-
duced the number of potential craftsmen. The men remaining in the villages
tend to be the unemployed, the elderly and the disabled. Through necessity
some of these men have become woodworkers, carving traditional objects for
sale as well as non-traditional objects to supply new demands deriving from
changing social and economic conditions.
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
Uses of wood and reed
Domestic equipment
1. Dishes
Wooden dishes, didelé (Fig. 48A), or platters made from soft woods such
as marula (Sclerocarya caffra) or mohoo (Ficus sycomorus) were traditionally
used for serving cakes of porridge.
The rough shape was carved with an adze and thereafter hollowed out and
finished off with a gouge, lehérd. The ridges made by this tool are clearly visible
on new dishes but they become smooth through use. The rim usually has a lip
and the base may be thickened or raised to prevent it wearing through from use
B
Fig. 48. A. Wooden dish, del6, SAM-—9675. Underside shows the marks left by the tool,
leh6r6. B. Calabash vessel and wooden imitation, SAM—9686 and SAM-9695. (All Krige
Collection, 1936-8.)
LOBEDU MATERIAL CULTURE 105
on the ground. Repeated washing and scouring with abrasive sandstone gives
used dishes a smooth finish.
Demand for wooden porridge dishes has declined but they are still oc-
casionally made and used. Many families use the calabash khethéba, made by
women, for the same purpose. An interesting wooden vessel in the Krige Collec-
tion is carved to resemble the shape of this type of calabash vessel (Fig. 48B).
2. Milk-pails
Milk did not form an important part of the Lobedu diet and the need for
milk-pails, dikhamél6é (Fig. 49), was, therefore, not great. During fieldwork in
Fig. 49. Herd-boy holding a milk-pail, khamélé. Photo: E. J. Krige,
1936-8.
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
the 1970s pails were not seen being made but, according to woodworkers who
had made them in the past, soft woods such as marula or wild-fig were most suit-
able as they did not easily crack. Other woods mentioned were mbidibidi
(Ekebergia capensis), morula moobyani (Lannea discolor) and mobeda-khamélé
(not identified).
Two types of milk-pail are represented in the Krige Collection. A cup-
shaped pail (Fig. 50) referred to as théhdé-ya-tshwene, literally ‘head of baboon’,
is the older form of pail and is similar to early Venda specimens (Van Warmelo,
in addendum to Curson 1932: 58). This type of pail was rarely seen in the 1930s
but the taller, narrower pails (Fig. 50) were still in use concurrently with pur-
chased buckets (Krige notes).
ans
Fig. 50. Two types of milk-pail: khamél6, SAM-9716; and théhé ya tshwene, SAM-9718.
(Both Krige Collection, 1936-8).
3. Mortars
Woods ranging from soft to very hard are used for mortars, mafudu
(Fig. 51). A soft wood has the advantage of being light to move and easy to
carve but has the disadvantage of wearing down and weathering quickly; hard
wood is difficult to work but more durable and resistant to termites. Woods used
by carvers interviewed in the 1970s included marula, motémé (Adina micro-
cephala) and mohoo. A carver often took his tools into the bush to work nearer
the source of the wood. Once the tree had been felled a block of suitable size
was cut from the trunk with a saw and was carved very roughly with an axe. The
mortar was shaped with an adze and gouge, and the hollow was chiselled out
LOBEDU MATERIAL CULTURE 107
Fig. 51. Woman using a wooden mortar and pestle for stamping maize. Note the winnowing-
baskets used during the stamping-process and the grain spread out to dry in the courtyard.
Molototsi valley, 1975.
with a long metal tool often made from part of a ploughshare. The finishing was
done with a small adze and /ehdr6 gouge. Finally a file or sandpaper was used
on the surface.
Where women still stamp maize by hand, mortars have remained in use and
the porridge made from this meal is much preferred to that made of bought
maize-meal. Furthermore, the different stages in stamping and winnowing pro-
duce meal ranging from a coarse to a fine texture, which gives variety to the
staple diet (see p. 78) and the whole-kernel grain is far more nutritious than the
refined product sold in the shops. None the less, pressure on land has resulted in
most families having to depend on bought maize-meal. The frequency of stamp-
ing maize at home is consequently much reduced.
Although ubiquitous in the 1930s, the free-standing mortar, Jefudu, was in-
troduced into the Lobedu area by Tsonga-speakers. Mortars were widely
adopted but until the 1950s were not allowed to be used inside the capital be-
cause of their Tsonga association (Mantwa Modjadji 1981 pers. comm.). During
the 1930s there was a special area outside the capital where women stamped
maize in these mortars (Fig. 52).
The older method was to stamp grain in a hole in the ground in which a hol-
lowed block of wood was set (Krige 1982: 16). The women worked in kneeling
position. This method was well suited to the stamping of sorghum and millet but
was not as effective for stamping hard maize kernels. The practical advantages
of the /efudu mortar for pounding maize seem to account for its widespread
adoption.
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 52. Women stamping maize in wooden mortars outside Modjadji’s village. Note the large
number of mortars in use, reflecting the supply of maize, and the use of winnowing-baskets.
Photo: E. J. Krige, 1936-8.
Fig. 53. Women using wooden pestles to thresh millet in the fields. The hollow in the ground
has potsherds embedded in the base to give it a hard surface. Photo: E. J. Krige, 1936-8.
LOBEDU MATERIAL CULTURE 109
4. Pestles and beaters
Long heavy pestles, mese, are used for stamping maize in a mortar and in
the 1930s were used for threshing grain in hollows in the ground (Fig. 53);
shorter pestles are used for stamping nuts in small mortars and for pounding clay
on a stone before use (see Fig. 4A). Woods specially favoured for this purpose
are mokwal6 (Acacia nigrescens and Acacia polyacantha), mothanari (Colophos-
permum mopane) and morididi (Combretum imberbe). In general a pestle is
made of a heavier wood than the mortar. A large club head is shaped at the
upper end to give it additional weight.
A lighter pestle, kherégélé, used for grinding tobacco into snuff (Fig. 54), is
a straight branch tapering slightly at the grinding-end. It is used with a clay pot
that has a striated inner surface (see Fig. 14 and p. 66).
A short-handled beater, khevhad6 (Fig. 55), with a flat working-surface is
used for compacting floors and ledges of huts and courtyards. Some potters use
this tool to shape the pot when it is leather-hard. A khevhado is usually made in
Fig. 54. Snuff being ground using a pestle, kherégélé, and a pot with inner
striations (see Fig. 14), Modjadji’s village, 1973.
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 55. Floor beater, khevhadé6, SAM-9726, Krige
Collection, 1936-8.
hard fine-grained woods, such as the Combretum species, so that it does not
wear down easily.
5. Spoons
Spoons, mafo, used for stirring and serving food are made from a variety of
fairly hard woods. Moataba (Pterocarpus rotundifolius, round-leaf kiaat), is
specially favoured as its branches are usually straight and thin; mosese (Pelto-
phorum africanum) and moila khipala (not identified) are also used but marula
and wild-fig are considered too soft to make durable spoons.
Lobedu spoons (Fig. 56) have almost flat bowls and they are not used as
eating-utensils or for handling liquids. The main use of the larger spoons is to
Fig. 56. Spoons and stirrer stored under eaves, Molototsi valley, 1975.
LOBEDU MATERIAL CULTURE east
stir porridge as it thickens and to serve the stiff porridge from the cooking-pot
into calabashes or wooden dishes in smooth, evenly rounded cakes. The rim
round the bowl of a spoon used for serving porridge prevents the hot porridge
from running over the sides and allows a smooth cake of porridge to be formed
(Krige 1981 pers. comm.). Spoons without rims are used mainly for stirring or
scraping down the sides of the cooking-pot. Smaller spoons are used for prepar-
ing relish in fairly small pots.
Wooden spoons are still made and they are in daily domestic use. Washing
with abrasive sandstone eventually gives the spoon a smooth, seasoned patina.
6. Stirrers
The traditional stirrer or twirler, lefhéth6 (Fig. 57), was made from
motomé (Adina microcephala) wood. Young motémé trees have a central
branch from which shoots radiate out in a whorl. The twirler was cut and
Fig. 57. Traditional stirrer, lefhéth6, SAM-9698, Krige
Collection, 1936-8.
trimmed from the growing tops of young trees, which gave the utensil its typical
form. A newer wire stirrer, lefhéth6 la darada, is now popular. This is made
from a straight rod of wood (usually moataba) with two wire rings attached at
one end (Fig. 56).
A lefthéthd is used in the early stages of cooking porridge to prevent lumps
forming when the meal is added gradually to a pot of boiling water. A woman
rotates the stirrer between her palms so that the porridge is mixed with a rapid
twirling action. When the porridge thickens she changes to the heavier /efo.
7. Stools and headrests
Stools, zwidulo, are made from any tree that has branches of appropriate
natural shape. In the khéré men sit on these seats while discussing village mat-
ters (Fig. 58A). In summer they are usually found under a large shady tree and
in winter or cold evenings round the fire-place in the khdr6d. Smaller stools,
112 ANNALS OF THE SOUTH AFRICAN MUSEUM
B
Fig. 58. A. Men sitting on wooden stools in the khéré of Mambeolo,
made ‘riempie’ chairs, Semosa’s homestead. 1973.
1975. B. Locally
LOBEDU MATERIAL CULTURE 113
carved from logs, are found in cooking-huts. Riempie chairs (Fig. 58B), based
on those seen in farmers’ homes, were made by some carvers in the 1930s and
were still made in the 1970s.
The most common headrest, kheseaméld, is simply a block of wood or a
bundle of old rags placed under the sleeping-mat. Two carved headrests
(Fig. 59)A—B) were collected in the Lobedu area in the 1930s but such headrests
Vz
Fig. 59. Headrests, zwiseamél6. A. SAM-9730.
B. SAM-9731, made by a Tsonga craftsman. (Both
Krige Collection, 1936-8.)
were not widely used. One is known to have been made by a Tsonga craftsman
and present-day Lobedu woodworkers confirmed that carving of this nature was
done mainly by Tsonga speakers.
8. Strainers
A wooden strainer for beer, sef6 ya hothéd6 vyalwa (Fig. 60A-B), is a
fairly recent addition to Lobedu household equipment. It can strain much larger
114 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 60. A. Sieve, sef6, made of a wooden frame with wire-gauze stretched across the base,
used for straining large quantities of beer; clay beer-pot in foreground. Modjadji’s village,
1975. B. Sieve in use at Modjadji’s village, 1975.
quantities of beer than the woven strainer, which it has largely replaced. An ex-
ample (SAM-10881), purchased in 1976 from a craftsman who brought these
strainers to sell at the market at the capital, is made of pine and blue-gum wood.
Pine is used for the sides of the box-funnel and gum for the lateral supports and
handle. The pieces of wood are joined with corrugated fasteners and nails, and
strips of metal are nailed over the corners and side joins to give them extra
strength. Wire-gauze and a lattice of pieces of hoop metal are nailed across the
base opening to form the sieve through which the beer is poured.
This type of strainer is a modern innovation related to an increase in the
brewing of beer for sale. In the 1930s the selling of beer was very uncommon
and beer was generally brewed on a relatively small scale, preferably from sorg-
hum or millet. The woven strainer (see p. 86) was used by wringing and squeez-
ing the beer into an open-mouthed pot, lebéda (see p. 66). Beer was consumed
at home and in a number of socially important contexts such as paying tribute,
showing hospitality or gratitude, and honouring ritual obligations. This pattern
of beer-drinking has continued to the present but, in addition, women now brew
LOBEDU MATERIAL CULTURE 1S
beer for sale as it is a way to earn money at home. Maize-meal is bought at the
store, often on credit against later payment when the beer is sold, brewed in
large tin drums, strained through a wire-mesh sefo into another large metal
drum, and served in glass jars or mugs of fixed volume. Thus the change in the
pattern of brewing is reflected in the utensils used in the preparation and con-
sumption of beer.
Hunting-equipment and weapons
Objects having wood or reed as a component are described here and refer-
red to in other sections where relevant.
1. Arrows
Arrows (Fig. 61A—G) are distinguished according to their type of head:
mosévhé wa phagane is an arrow with a flat, unbarbed metal head; mosévhé wa
mogovho is an arrow with a long, straight metal head; mosévhé wa digoébe (lit-
erally a fishing-arrow) is an arrow with a barbed metal head; and thebola is an
arrow with a knobbed wooden head (Fig. 61F-—G).
Arrow shafts were generally made of reed into which the tang of the head
was fitted and the join bound with sinew, bast or stripped creeper. A hard wood
such as mosese (Peltophorum africanum) was used for the knobbed head of the
thebola, which was used in hunting birds and small buck.
2. Bows
Formerly the bow, vhora, was the principal weapon in warfare and in hunt-
ing. Both these activities had ceased by the 1930s.
Mofwata (Brachylaena transvaalensis), which was considered a particularly
good wood for making bows (Krige notes), is described by Palmer & Pitman
(1972: 2149) as ‘strong, elastic and durable’—ideal qualities for the purpose.
Examples in the Krige Collection have staves, rounded in section, which taper
from the grip to the slightly shouldered ends. The string of twisted thong is
fastened to each end with a knot (Fig. 62).
By rubbing the stave with fat it was kept pliable, and in cases when repair
was necessary the break or crack was bound with wire and covered with a piece
of cow-hide (SAM-—9836, Krige Collection).
3. Clubs
Wooden clubs were not part of Lobedu material culture. The term théga, a
club, is a Lobedu rendering of the Tsonga term nhonga, and the weapon was
never widely adopted by the Lobedu (Krige 1983 pers. comm.).
4. Hafts of tools and weapons
Weapons and tools used for a variety of purposes were hafted in wood.
Fairly hard woods were selected for this use as it was important that the wood
should not crack easily. Moataba (Pterocarpus rotundifolius) and mordéd6é
116
ANNALS OF THE SOUTH AFRICAN MUSEUM
Wy BLA
[
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cm
Fig. 61. Types of arrows. A. Unbarbed arrow, mosévhé wa phagane.
B, E. Barbed arrows, mosévhé wa digdbe. C, D. Long-tanged arrows,
mosévhé wa mogovhd. F. Wooden-headed arrow, thebola. G. Butt end
of the thebola.
LOBEDU MATERIAL CULTURE LT;
Fig. 62. Detail of bow-string attachment.
(Pterocarpus angolensis) were often used but many other woods were also suit-
able. The handle, mofhenye, was carved to the required shape and a socket for
the attachment of the head or blade was burnt through the haft with a hot
poker, moro (see Fig. 110A). When a man needed a new tool or weapon he
would commission a specialist smith for the head and make the haft himself.
Hafts were generally undecorated but usually well proportioned in relation
to the head and well balanced for handling. Occasionally a knife, mofhaga, had
a band of decorative carving on the hilt (see Fig. 111A).
> fraps
A cane-rat trap, mothéma (Fig. 63A), was made from a branch of moloha
(Acacia ataxacantha) split into a number of sections for most of its length. The
thicker end was left whole and the split sections were spread apart and fixed in
position by being tied to graded wooden rings (made of pliable wood or creeper
stem). The overall shape was that of a funnel. String was wrapped round the
sticks and passed from one to the other to fill in the gaps that were large enough
for a rat to pass through. Herd-boys placed the trap in the grass where rats,
magwelele, were known to run, beat the bush to scare the rats into the trap and
then killed them with a knife or a small spear.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 63. Traps for cane rats. A. Mothéma, SAM-9862, Krige Collection, 1936-8.
B. Wire mothéma, Modjadji’s village, 1973.
A modern version of the mothéma is made in wire (Fig. 63B). The opening
of the trap has a flap that opens inward only so that the animal gets caught in-
side and cannot escape.
A trap, khethédélé (Fig. 64), for birds consisted of a ring of wood (a pliable
stick bent into a circle and tied with bark) with a mesh of bark strips tied across
it. Slip-knots made of the tail hair of an ox were attached to the cross-strips and
the trap was covered with a thin layer of soil so that only the knots of hair were
LOBEDU MATERIAL CULTURE 119
Fig. 64. Khethédélé, trap with slip-knots for catching birds,
SAM-9863, Krige Collection, 1936-8.
visible above ground. Grain or seed sprinkled between the knots attracted birds
and their legs were caught in the slip-knots.
The mothéma and khethédélé, as well as a number of other traps that in-
volved the use of stones and string or pliable wood under tension, were made by
herd-boys who spent much time in the veld looking for edible wild fruits or
honey and making traps for birds and small animals. They were much less com-
mon in the 1970s than in the 1930s.
The trap for termites made in the side of a termite-heap using sticks and a
clay pot was described in an earlier section (see p. 68).
Musical instruments
1. Drums
Drums played a significant role in public life and were played on many oc-
casions of ritual and social importance. Drums were never individually owned.
Every district head owned a set of drums and was responsible for their upkeep.
When a new district head was appointed the chief handed over a drumstick as a
public sign of his office (Krige & Krige 1943: 126).
Lobedu drums for secular use are of two types: géma (Fig. 65A), a large
drum with hemispherical resonator, played with a drumstick; and thithimedzh6é
(Fig. 65B), a long, cylindrical drum made in various sizes, played with the palms
of the hands. Drums are made by specialists who take orders and receive pay-
ment for their work. In the 1930s in the Venda area the cost of a large drum of
similar type to the Lobedu gdéma, was an ox or its equivalent (Kirby 1934: 36).
120 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 65. A. Large géma and small thithimedzhé drums at headman Mohale’s village, 1975.
B. Large thithimedzhé drums on the verandah of a sleeping-hut in the special part of the
capital used by Modjadji and her wives, 1976.
LOBEDU MATERIAL CULTURE 121
The cost in 1976 was in the order of R20 to R30 but less if the wood was sup-
plied by the buyer. For making drums soft-wooded trees such as mohoo (Ficus
sycomorus), movhamba ngoma (Albizia gummifera) and marula (Sclerocarya
caffra) are used. The carving-tools are adzes and gouges of various sizes, and a
long, flat piece of sharpened scrap metal is used as a chisel.
A goma drum is made in the following way: the resonator is carved from a
solid section of tree trunk and worked so that the long grain of the wood runs
laterally across the drum and not vertically down its length. The resonator is hol-
lowed out until the walls are about 2—3 cm thick, and a small hole is made in the
centre of the base, which is thicker than the sides. A band of relief carving
usually encircles the bowl at its widest diameter. The goma in the Krige Collec-
tion has handles resembling interlaced loops carved at the four ‘corners’ and a
band of carving between the handles. These features are characteristic of many
drums from the Lobedu area and are also typical of the Venda ngéma described
in detail by Kirby (1934: 34-38). Holes round the mouth of the resonator are
burnt with a hot poker and a piece of wet cow-hide is stretched over the head
and pegged into position through the holes. The hair is removed from a circular
patch in the centre of the skin. An additional strip of hide with slits made in
Fig. 66. Drums being played by women at a gdésha dance. The men on the right are playing
reed pipes (see Fig. 70). Photo: E. J. Krige, 1936-8.
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
position of the pegs is fastened round the rim to strengthen the attachment. Be-
fore the wet skin is stretched over the resonator a few river stones are placed in-
side the drum. According to Krige & Krige (1943: 221) the reason for this may
be related to the ‘cooling’ quality of the stones. Coolness is perceived as a posi-
tive quality which can counteract negative forces. It is also likely that the
conventional attributes of the gOma and the manner of beating it have special
meanings and symbolic associations in specific social contexts and that many of
these are related to chieftainship and to fertility.
Two or three cylindrical drums of different sizes are played together with
the gdma. The longest and most important of these is gaedis6, which sounds the
entry of solo dancers; the other two are both called thithimedzho. All are played
by women using the palms of their hands. They are not played in upright posi-
tion but are placed on their sides and tilted upwards when being played
(Figs 66-67).
The construction method for both gaedis6 and thithimedzho is the same.
The wood said to be best for these drums is mordd6é (Pterocarpus angolensis)
but softer woods are also used (Jan Lebiya 1976 pers. comm.). The resonator is
carved from a solid block of wood, the length of resonator foilowing the grain.
The tools used are the traditional adzes and gouges as well as a piece of sharp-
ened scrap-iron used to chisel out the shell. Some, but not all of these drums,
Fig. 67. Women playing drums at a Saturday beer-drink near Molabisana’s shop, 1973. Note
the tin g6ma on the right.
LOBEDU MATERIAL CULTURE 123
have a handle on one side. Holes round the mouth of the resonator are made
with a hot poker, and pegs for attaching the skin are carved in a hard wood. As
for the goma drum a piece of wet cow-hide is stretched over the head and verti-
cal slits are made to coincide with the positions of the holes for the pegs. A band
of hide with horizontal slits in the position of the pegs is wrapped around the
head and the pegs hammered into the rim through the slits in the two layers of
skin. These drums are seldom decorated.
A frame-drum, khethadada (Fig. 68), played during maldb6 spirit-possession
rituals is made in the following way. A piece of wet goatskin, with the hair re-
moved, is stretched over a broad hoop of pliable wood or metal (for example
the frame of a bicycle wheel) and the overlapping edges of skin are cut into
strips, twisted and tied on the underside to fasten the head in position and form
the grip for the player to hold. Two or more khethadada ‘tambourines’, beaten
with sticks, are played together with tshélé and mathodzi rattles (see p. 153) and
other drums to accompany the dancing of a possessed person and to encourage
the spirit to manifest itself (Krige & Krige 1943: 243). The Tsonga cult of
Fig. 68. Frame drum, khethadada, and drumstick used to accom-
pany maldbé dancing. Note attachment of skin on underside.
SAM-9827, Krige Collection, 1936-8.
124 ANNALS OF THE SOUTH AFRICAN MUSEUM
possession from which Lobedu maldbo practices derive has been described by
Junod (1927: 479-504) and the associated musical instruments by Kirby (1934:
41-44). The cult among the Kgaga has recently been described by Hammond-
Tooke (1981: 104-111).
In addition to the drums for secular use there are four sacred drums (dig6-
mana) that are held in great awe and are associated with the chief and the wel-
fare of the people. The following information was recorded by Krige & Krige
(1943: 126-127). The sacred drums are similar in shape to the ordinary g6ma
drum but they are played by hand instead of with a drumstick and their sound is
believed to please the ancestral spirits. The largest of the four, phataje, stands
about 120 cm high, next in size is tanga, then pekahare, and rangwedi, the
smallest but most important. All four drums are said to contain a human skull
instead of the stone that is usually put inside a drum. In addition, rangwedi has
strips of human facial skin placed under the ox-hide head and is smeared on the
outside with khetshila, ‘dirt’ from the corpse of a person of noble blood who was
killed for this purpose. The maintenance of the sacred drums, which involves
treating them with fat and ochre to protect them from infestation by insects, the
renewal of the skins, and the repair of the hut in which they are kept is the
special task of the Rabothada family, which has historical links with Phalaborwa
where the institution of sacred drums was strongly developed (Krige & Krige
1943: 127).
The sacred drums were played only on certain occasions of ritual import-
ance, for example the gOmana ceremonies, and unlike the ordinary drums they
were always played by men. Their beating was associated with asking the ances-
tors for rain and with the seasonal cycle. So important were these drums that
they were looked upon as having supernatural power (Krige & Krige 1943: 127).
Although they are still kept in good repair, the sacred drums of the royal family
have not been beaten regularly since the last decade of the nineteenth century
and not at all since before the 1930s. The reason for this is said to be that killing
for ritual purposes has long been prohibited by law and the necessary parts of a
human body are thus no longer obtainable (Krige 1983 pers. comm.).
2. Stringed instruments
Kirby (1934: 196) grouped southern African stringed instruments into three
general categories according to the fundamental tones produced and the har-
monic sounds generated by these fundamentals. The Lobedu stringed instru-
ments khekhaba, khedzhél6 and kheddlédélé (Fig. 69A—-B) fall into Kirby’s
groups one, two, and three respectively.
The khekhaba is a musical bow in which the fundamental tone of the string
and its harmonics are sounded as a chord. As the string is tied back near the
centre it yields two chords, depending on which part of the string is struck. The
bow is made of a well-seasoned branch with the bark removed. It is fitted with a
string (of wire, twisted sinew or hair) tied back by a loop of wire, sinew or fibre
near the centre. At this point a calabash resonator is attached to the bow. The
LOBEDU MATERIAL CULTURE 125
Fig. 69. Stringed instruments. A. Khedzhdl6, SAM-9819. B. Kheddléddél6, SAM-9821.
(Both Krige Collection, 1936-8.)
string is struck with a thin stick or reed. It is usually played by unmarried men
on its Own as an accompaniment to a song (Kirby 1934: 207).
The khedzholo (Fig. 69A) is an instrument that is played so that certain
harmonics of the string are isolated to produce a melody. The stave is usually
carved from a branch of soft wood about 80—90 cm long and 6 cm in diameter.
The bark is removed, a flat surface formed along one side of the wood, and the
pith hollowed out, leaving one end solid. Through this end a hole is bored and a
tapered peg fitted. A wire is attached to the upper end of the tuning-peg and to
the opposite end of the wooden stave or sounding-board. A notch on the stave
makes it easier to hold the khedzholé steady when in use. A small friction-bow
for playing the instrument consists of a stick about 10 cm long with a string of
tail hairs or plant fibre attached to notches at the ends of the stick.
The player holds the khedzhdl6 with the open end resting on his left
shoulder. A calabash or tin resonator is often placed over the end to increase
the sound produced by stroking the wire with the bow held in the right hand,
and varying the tension of the string with the left hand. Only men play this in-
strument, either alone or to accompany a song.
The khed6éléd6l6 (Fig. 69B) is a musical bow in which the harmonics of the
string are used in conjunction with their fundamentals to produce the melody.
The bow stave, which is always thicker in the centre than at the ends, is con-
structed in one of two ways—it may be carved as a whole from a solid branch of
flexible wood or the central part and the ends may be carved separately and then
fitted together. In the latter case it is usual for two different kinds of wood to be
used—a fairly soft wood for the thick section and pliable wood for the ends. In
both cases the ends are bent to the required curve while green and allowed to
dry in that position.
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
The string is tied to notches cut at the tips of the stave and a piece of cotton
or sinew is attached near the centre of the string to draw it back towards the
stave. It is wound round the stave to secure it in position. The mouth, placed at
one end of the thick part of the stave, acts as a resonator and the string is
plucked in various ways with a plectrum (a small stick or a thorn) to produce a
number of tunes, often wistful or sad. It is played only by men or boys and
sometimes the player sings an accompaniment. Formerly, if a young unmarried
man played this instrument all night people knew he was longing for a wife
(Krige notes).
3. Wind instruments
The playing of the reed pipe ensemble, mothavha, accompanied by drum-
ming and the performance of the gésha dance, was an important form of social
activity and musical entertainment among the Lobedu. Although by the 1970s
_ the inter-district gésha dances associated with individual reciprocities had disap-
peared, those associated with dancing for rain at the capital or at the end of a
period of mourning continued, albeit less frequently than in the 1930s (Krige
1982: 23). The number of reed pipes in a set can vary from more than twenty to
less than ten. Each man plays a single pipe that has a fixed pitch and produces a
single note, the longest pipe giving the deepest note. Each pipe has its own
name denoting the part it plays within the ensemble and how it combines with
other members of the set.
A set of twenty-two reed pipes (SAM-—9812) collected in the 1930s is named
and graded as follows (Krige notes):
1. Ledzekha la pele—the longest pipe.
2. Lembo la dzusi la giba—played after pipe 14 to introduce a song, giba.
3. Ledzekha la gamorao—played with 1.
4. Ledzekha—played with 1 and 3.
5. Lembo la dzusi la vesa—played with 2.
6. Lembo—played with 2 and 5.
7. Ya ga ntau ha thakhudi—played after 12.
8. Lembo—played with 2, 5 and 6.
9. Moholo a khomo.
10. Khomo—played with 9.
11. Khomo.
12. Thakhudi.
13 2Giba:
14. Dzusi ya giba—awakens or heralds 13, starts the giba song.
15. Tateledi.
16. Dzusi.
17. Dzusi ya vesa—introduces the vesa song.
18. Pegudi.
19. Thezwane.
LOBEDU MATERIAL CULTURE 127
20. Ya ga ntau a thezwane—played after 19.
21. Thezwane.
22. Khenthé kha dzusi a giba—high-pitched pipe for ‘rousing’ or introduc-
ing 13.
Some of the pipes (Fig. 70) are made of cultivated reeds, mothékhathébye,
which are stronger than the indigenous mbaseriti reeds formerly used for reed
(\
aU
XX
(\
UvAvaNal
om
LOOK,
A 2
PS
Fig. 70. Some examples of the pipes in a set of twenty-two reed pipes, mothavha,
SAM-9812, Krige Collection, 1936-8.
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
pipes. Not all pipes in the set are made of this type of reed; nine are of a thinner
reed, not identified. Lobedu reed pipes are made in the same way as the Venda
pipes described in detail by Kirby (1934: 162). The reeds are cut to the required
length so that a node forms a natural plug at the lower end of the pipe. To bore
through intervening nodes a length of red-hot iron is used. If ornamented, the
pipes are incised with a sharp knife. The pipes are tuned against each other and
can be cut down to raise the pitch. A complete set of pipes is usually kept in a
special skin bag, saga ya mothavha, and many of the pipes bear some distin-
guishing mark so that the player can easily recognize the pipe that he usually
plays.
Reed pipes are played only by men, who dance in a circle while playing,
causing the melodic illusion of changing sound sequences that would not occur
were the players to remain stationary. The dance-master, malogwani, signals the
start and changes in the pattern of the dance but in actual performance leader-
_ ship changes continually as one dancer after the other takes the centre position
in the circle of dancers. An accompaniment of drumming is played by women on
a goma and three thithimedzo drums (see Fig. 66).
In addition to the reed-pipe ensemble, there are a number of flutes and
whistles made of bamboo or wood that are played on their own, especially by
herd-boys. The term ‘flute’ is used here for pipes that can produce more than
one note, and ‘whistle’ for those that produce only a single note.
The khethodigoé (Fig. 71A) is a transverse flute made of indigenous mbase-
riti reed closed naturally at both ends by nodes in the reed. The embouchure
near one end and three finger-holes near the other end are bored through the
reed with a red-hot poker. A number of tunes can be played on this flute. The
rodigo flute (Fig. 71B) is made from a length of mbaseriti reed open at both
ends. The player blows over two V-shaped notches at one end and varies the
tune by fingering the other end. The /engwale flute (Fig. 71C), made of wood, is
played in the same way. The example in the Krige Collection is made of moro-
gdl6 wood (Carissa edulis) a fine-grained wood that does not split easily. A
whistle, naga, made of horn, reed, or wood, is a pipe closed at one end and
sounded by blowing across the open end. A naga (Fig. 71D) collected in the
1930s is made of wood shaped to resemble a horn and decorated with wire bind-
ing. A feather kept in the open end is used as a pipe cleaner.
A small vibrating wind instrument that has great ritual importance among
the Lobedu is the mantsaakhéda (Fig. 71E), which consists of two pieces of
wood wider at one end than the other and slightly concave on the inner surfaces,
between which is inserted a piece of bark or any material that will vibrate when
blown. Rubber was used in a specimen collected in 1970. The mantsaakhéda is
surrounded by secrecy and is used by old men who impersonate the voices of the
ancestral spirits, zwidajane, in certain religious ceremonies, for example the thu-
gula harvest thanksgiving and the khetshila purification ceremony after the death
of royalty. The spirits are believed to be hidden in the near-by bush and to ex-
press themselves by whistling. Messages are conveyed by varying the tone of the
LOBEDU MATERIAL CULTURE 129
20
Fig. 71. Wind instruments. A. Flute, khethodig6, SAM-9814. B. Flute, rodig6, SAM-9815S.
C. Flute, Jengwale, SAM-9816. D. Whistle, naga, SAM-9817. E. Whistle, mantsaakhdda,
SAM-9824. (All Krige Collection, 1936-8.)
whistling, which can be translated into the tone patterns of the language. The
mphenyi instrument played by khiudogane in the vyali school (see p. 99) oper-
ates on the same principle. The vibration of the membrane across the end of the
reed pipe produces the strange eerie sounds by which khiudogane communi-
cates.
Wind instruments made of horn are described in a later section (see
p. 166). They include whistles and the horn trumpet, phalafhala, which is
associated with authority and is blown to signal important announcements.
4. Idiophones
In the 1930s xylophones, dibela, with calabash resonators (see p. 153) were
used in the Lobedu area by itinerant minstrels who played the xylophone and
sang humorous songs in return for food and lodging (Krige & Krige 1943: 316).
There is no record of xylophones being made by local craftsmen and it is prob-
able that the instruments used were of Venda origin (Kirby 1934: 56).
Also in use in the 1930s was a plucked instrument, deze, consisting of a flat
wooden resonator set under a series of iron tongues that were plucked with the
thumbs to produce the sound. By adjusting the length of the iron tongues the in-
strument could be tuned. This sansa-type instrument was played mainly by
young men (Krige 1982: 22).
The wooden clappers mewasa (Fig. 72) used by vyali girls also fall into this
category of instruments made of inherently resonant materials.
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 72. Wooden clappers, mewasa, used by girls during the vyali initiation, SAM—9858, Krige
Collection, 1936-8.
Playthings
In making playthings, children acquired a lot of useful knowledge of the en-
vironment and an early training in craftwork. Herd-boys made everything they
used in hunting and in playing games. They got to know the qualities of the dif-
ferent woods and their toys changed with the seasonal availability of the natural
materials, grasses (see p. 89), fruit shells, or maize stalks. Wooden toys in-
cluded wagons, motor cars, guns, sledges, and clubs for playing khdr6r6, a
game similar to hockey (Krige & Krige 1943: 108). Today boys still make their
own toys but the raw materials seldom come from the veld. Wire, bottle-caps
and tin-cans are used most effectively for making cars, tractors or lorries.
Ritual objects, figures and carved poles
At present little is known about the traditions of figurative carving among
the people of the northern Transvaal, although a thesis based on field research
undertaken in the 1970s is in preparation (A. Nettleton, Department of Fine
Art, University of the Witwatersrand, 1981 pers. comm.).
Carved figures of symbolic significance in initiation rituals are used by the
Venda (Stayt 1931: pl. 31) and have also been collected among the Kgaga of
Maake and the Narene of Sek6r6r6 (Potchefstroom University Collection). A
male and a female figure in the collection of J. Witt of Tzaneen were said to
have been among the Lobedu digéma revealed to initiates during a vyali
initiation (see p. 99). At the end of the vyali the costumes and all associated
objects should be burnt, so it is unusual for them to be represented in museum
or private collections. Furthermore, the secret nature of these objects forbids
their description by those who have seen them. A géma is essentially something
131
LOBEDU MATERIAL CULTURE
1973»
the capital,
’s village,
dji
.
ja
6rd of Mod
oro surrounded by forked poles.
B. The kh
Fig. 73. A. Layout of huts around central kh
——————— ES
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
secret and mysterious but it may be intrinsically very simple. The ritual context
of its revelation imbues a géma with its awesome quality.
The wooden clappers, mewasa (Fig. 72), used by vyali girls to accompany
their songs, are simply two flat pieces of wood carved to a point at one end and
a handle at the other. No significance is attached to their manufacture but, once
used, they acquire significance and they must be burnt at the end of the school.
For the vyali initiation a row of debarked poles is set up in the khdéro in a
specially medicated furrow at the back of a carefully prepared platform, /e-
vhalélé la vyali, on which the initiates stand when singing. In front of this is the
kholon6ni shrine, a circular mound of clay in which a river stone is embedded
and in the centre of which is a forked, medicated pole of musoso wood (Termin-
alia sericea, vaalboom) believed to have protective powers (Krige & Krige 1943:
136).
The khéro of the Queen’s village (Fig. 73A) is surrounded by a palisade of
pointed or forked poles (Fig. 73B) that may only be used in the villages of chiefs
or headmen. Headmen from all the districts are called up to provide poles for
the Queen’s kh6ré when it is renewed. In this way the khdro of the capital ex-
presses the solidarity of the chiefdom. The entrance to the khdro is also the en-
trance to the village and, as such, it is protected with posts of special wood
(kherale or moludu) treated with medicines when they were set up. Beneath the
threshold a medicated wand or a river stone is buried to neutralize or ‘cool’ any
evil that may threaten the village. These measures are but a very small part of a
complex scheme for protection of the village.
Fig. 74. Carved khéro poles at the capital, 1973.
LOBEDU MATERIAL CULTURE 133
Some of the khéro poles are carved in the form of stylized figures (Figs 74,
75A-C). Their meaning is not fully understood. According to Krige (notes) they
were made to honour the Queen. Schlosser (1975: 95) stated that they were ex-
clusive to the Queen’s khoro but had no other significance. According to J. Witt
of Tzaneen who has made a collection of carved poles from Modjadji’s khéré,
craftsmen of particular skill carved the poles to distinguish their contribution to
the khéro.
Fig. 75. Carved khér6é poles. A-B. From the capital, now in private collection of J. Witt.
C. Collected when the poles of the khér6 at the capital were being renewed, SAM-9751, Krige
Collection, 1936-8.
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
Structures
Although the study of settlement pattern, hut forms and _building-
techniques was not a main focus in this study, structures and building-techniques
recorded in the 1970s are briefly described and supplemented by data and photo-
graphs from the 1930s provided by E. J. Krige.
The smallest cluster of huts within a village consists of a sleeping-hut, ndo,
a cooking-hut, khetaga, and a courtyard, mosha, formed by the space between
them. At the capital these pairs of huts are arranged in a circle round a central
courtyard, the khdér6é (Fig. 73A). Sleeping-huts all face the khdér6é and cooking-
huts have their backs to the khdér6. Each wife in a polygynous family has her
own mosha, her own utensils, storage area, granaries, and fields. The household
of the chief wife of the head of the village is usually positioned opposite the
main entrance but, apart from this, there is no ranking of household units within
the village. In smaller settlements there is no khoro and huts are arranged in an
arc or straight line.
Fig. 76. Framework of a hut under construction in the 1930s. Photo: E. J. Krige, 1936-8.
LOBEDU MATERIAL CULTURE 135
i Auts
The oldest huts were built using the pole-and-daga method but even before
the 1930s sun-dried mud-brick construction was replacing the older method that
drew heavily on timber resources. Between 80 and 100 poles were needed for a
single hut (Fig. 76). The following description of the pole-and-daga method was
recorded in 1976 from wood-carver F. Semosa, and examples of the materials
were collected for reconstructing a hut at the South African Museum. Hard, ter-
mite-resistant woods such as moathaba (Pterocarpus rotundifolius), mosese (Pelt-
ophorum africanum), or moréjé (Dicrostachys cinerea) were said to have been
preferred, but many other hard woods were also used. It was often necessary to
straighten the poles before use. This was done ingeniously by making a series of
oblique cuts on the inner curve of the pole while it was still green and then
wedging it in the fork of a tree and bending it straight (Fig. 77A). In 1976 poles
straightened in this way were still visible in one of the oldest huts at the capital
(Fig. 77B). The process, -vhaga, was also used by the Venda (Van Warmelo
1937: 299; Van der Waal 1977: 79) whose hut-building techniques were similar
in many respects to those of the Lobedu.
The straightened poles were carved roughly to a point at one end and
embedded in the ground to a depth of about 40-50 cm, following a circle that
had been traced out on cleared ground. The soil was saturated with water to
facilitate embedding the poles. Flexible withies from trees such as mofwara
tsweni or mothala maligana (both Grewia spp.) encircled the poles horizontally
at regular intervals and were lashed on to the framework with strong bark fibre.
Both the laths and the bark had to be softened in water before use. The ridges
formed by the laths were usually accentuated in the plastering-process and made
into ledges in the walls. A mixture of mud and dung was applied by hand to the
framework both inside and out. This part of the work was done by women and
they also helped with transporting the materials from the bush and collected the
thatching-grass for the roof (Frans Semosa 1976, field records).
The conical roof structure of the oldest huts was made of closely packed
rafters (Fig. 78B). To construct the roof a central post, mosima, was erected in
the middle of the hut, and a working-platform was made by placing planks
across the walls and securing them to the central post. The pointed wooden roof
pinnacle, /Jenodu, was supported by the mosima while the rafters, levhalélé la
thaga, sharpened at the upper end, were tied to the pinnacle. The lower ends of
the rafters rested on and were tied to the poles forming the walls, /evhaléldé la
tshite. Flexible cross-laths tied to the rafters encircled the roof at intervals.
When the roof was complete the mosima was cut away (Krige 1982: 13).
The roof was thatched with bundles of grass placed on top of the rafters
(see p. 93). Krige recorded that in the 1930s the plastering of some sleeping-
huts continued up into the roof to form a plastered dome (Fig. 78A). “This dome
has circular rims marking the cross-laths which, beneath the plaster, hold to-
gether the framework of closely packed rafters’ (Krige & Krige 1943: 21). This
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 77. A. Straightening a pole by wedging it between two trees after oblique cuts have been
made on the curve of the pole, Semosa’s homestead, 1976. B. Hut poles showing the cuts,
Modjadji’s village, 1976.
LOBEDU MATERIAL CULTURE 137
Fig. 78. A. Plastered roof of a sleeping-hut. Photo: E. J. Krige, 1936-8. B. Traditional
roof structure of a cooking-hut. Closely packed rafters support the thatch. Modjadji’s
village, 1976.
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
feature, also found in Venda huts (Van der Waal 1977: 88), gave a very neat
finish to the hut interior. These architectural details disappeared when new con-
struction techniques were adopted. The plastered ridges were not made on mud-
brick walls, and ceilings could not be plastered when widely spaced rafters were
used (Fig. 79A).
Wooden doors used in sleeping-huts were carved from solid pieces of soft
wood such as wild-fig and hinged by means of projections on the upper and
lower ends that fitted into sockets in the lintel and threshold beams. A reed door
(see p. 85 and Fig. 16) was used in the cooking-hut.
By the 1930s brick construction, which had been taught by the mission since
the late nineteenth century, was replacing the older pole-and-daga construction
method, and legislation to protect many of the indigenous trees accelerated the
process by reducing the availability of wood (Krige 1941: 18). Bricks were made
by women and men using wooden box moulds (Fig. 79B) or tins and at the capi-
tal bricks were commissioned for 10 shillings per 800—1 000 bricks (Krige 1941:
18). The increased participation by women in hut-building compensated to some
extent for the absence of men working in towns, and building with sun-dried
mud-bricks has continued to the present both for the conventional round huts
described below and for more modern houses.
The ground is cleared and a circle drawn with a stick attached to a string
fixed to a peg in the centre of the circle. A shallow trench of stones is usually
laid as a foundation for the brick coursing. Mud plaster is applied between the
bricks and, when the wall is complete, to the interior and exterior. The initial
plastering is done by hand and a wooden tool, khevhado, is used for compacting
the plaster. The same tool is used at a later stage when the roof is complete for
compacting the mud floors of huts and courtyards, which are smoothed, -ridéla,
with a stone, thidél6, then smeared with dung, and sometimes decorated with
patterns made by running the fingers over the wet dung.
The traditional roof structure of closely packed rafters, which supported the
thatching grass without the need for sewing, was replaced by the ‘rondavel’ roof
of widely spaced rafters, usually of purchased blue-gum timber, on to which the
thatch was sewn. In the 1930s modern window and door-frames were rare except
at the Christian settlement, but by 1940 a few rectangular houses with windows
had been built at the capital (Krige 1982 pers. comm.). Subsequently many rect-
angular-shaped huts with thatch or tin roofs were built but in the 1970s, at the
capital and in smaller settlements, the circular-plan hut remained the most
widely used dwelling.
In the 1970s the apparent trend among non-traditionalists who could afford
the materials and were living within access to transport was toward modern
housing. Many people, however, still lived long distances from main roads and
the most frequently seen dwellings were mud-brick huts, which are low-cost
structures appropriately designed for a hot climate and a living-pattern in which
most activities take place out of doors.
LOBEDU MATERIAL CULTURE
Fig. 79. A. Modern ‘rondavel’ type roof structure with widely spaced rafters, plastered
mud-brick walls. Modjadji’s village, 1973. B. Mud-brick and mould, Ramatswalela’s
homestead, 1975.
139
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
2. Granaries
In the 1930s granaries were very much a part of Lobedu domestic life. Per-
manent granaries were built in the storage area, marala, behind the huts and
temporary granaries were built in the fields at harvest season. By the late 1970s
there was no need for granaries as there was not enough grain to be stored and
most families depended on bought maize-meal for their basic subsistence
requirements. Very few granaries were seen in use during fieldwork and the
traditional grain pit for maize had fallen completely into disuse. The following
descriptions are based on photographs and information recorded in the late
1930s by E. J. Krige (1982: 14-15).
Traditionally decobbed maize was stored for long periods in a grain pit,
kheledese (Fig. 80), usually situated in the cattle kraal. At the royal village the
pits were in the khér6. The pit was a large hole in the ground lined with dung
and grass; it had a fairly narrow opening but broadened out below ground level.
Before being placed in the pit the grain had to be completely dry to minimize
- = : . a
> -...
Fig. 80. Kheledese, pit for storing maize. Note depth of earth above opening of pit.
Photo: E. J. Krige, 1936-8.
141
LOBEDU MATERIAL CULTURE
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=e
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In
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it
6, granary formerly used for stor
Fig. 81. A. Disused kheash
of cycads, 1976. B. Kheashé w
have been adopted from Tsonga
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
the chances of it rotting. A stone was placed over the opening, which was then
sealed with dung and covered with earth to ground level. A stone was placed to
mark the stituation of the pit. Storage in the sealed pit changed the flavour of
the maize, which was then called madoni. Certain pits were known to produce
particularly good madoni and were used repeatedly.
The kheashoé (Fig. 81A) was a structure of poles, mabala, encircled by
cross-laths at the top, middle and base, and secured with bark fibre. The base
was raised about 20 cm off the ground on forked posts or large stones. This was
the traditional way of storing maize on the cob while drying. A similar structure,
also referred to as kheashé (Fig. 81B), but with grass woven between the poles,
was introduced by Tsonga immigrants. The grass was said to protect the grain
from being pecked at by fowls. This protection was only necessary after the
Tsonga practice of keeping fowls had gained acceptance among the Lobedu.
The Jedulu (Fig. 82) was a granary of roughly the same shape as the
Fig. 82. Ledulu granary for millet and sorghum, made
of a wooden framework plastered with clay. The
thatched roof must be removed to reach the contents.
Photo: E. J. Krige, 1936-8.
LOBEDU MATERIAL CULTURE 143
kheasho but the walls were plastered with daga and it had a conical thatched
roof that had to be lifted to reach the contents. It could be sealed for long-term
storage. The /edulu was the traditional Tsonga granary for decobbed mealies but
was widely used by the Lobedu and the Venda for sorghum and millet as well as
for the short-term storage of maize.
The /etole (Fig. 83) was a temporary granary erected in the fields for storing
sorghum before it was threshed. Structurally it was an inverted roof supported
by forked upright posts. It was lined and covered with grass to protect the grain
Fig. 83. Lejéle granary in which sorghum was stored in the fields before threshing.
Photo: E. J. Krige, 1936-8.
. from birds. A raised platform moak6 (Fig. 84), was also used in the fields for
temporary storage of millet and sorghum prior to threshing and winnowing. In
spring these platforms were used to stand on when scaring birds from crops. A
smaller permanent platform, vhoald, built inside the cooking-hut, was used for
keeping bags of grain off the ground to protect them from termites and other
insects. In the 1970s these platforms were still seen in some traditional house-
holds.
STONE
Oral traditions record a time in Lobedu history (probably in the late eight-
eenth century) when stone enclosures were built. The description (Krige &
Krige 1943: 7) of high stone walls, upright stone projections, and a stone seat for
the chief clearly link the Lobedu with the Venda and their Karanga ancestors
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 84. Moako platform used for temporary storage of sorghum in the fields prior to
threshing. Photo: E. J. Krige, 1936-8.
north of the Limpopo River. The ruins of these stone structures associated with
the reign of Chief Kheale are still standing, although much overgrown. The site
is strategically situated on a hill-top commanding a good view of the surrounding
country. In other areas there are remains of stone-walled enclosures for live-
stock and terraces of former occupation sites (Fig. 85).
In some villages stone-walled enclosures for livestock are still found and
stone is still used for the contour walling on the cultivated hillslopes. Many old
contour walls, maraka, are now overgrown but they show up well on aerial
photographs. Valley cultivation was at first practised only by Tsonga-speaking
immigrants but later shortage of arable land on the hillsides and the introduction
of an effective plough led increasing numbers of Lobedu to cultivate the flatter .
ground. The plough, however, destroyed the protective verges of the fields and
broke down many of the stone contour walls that prevented erosion (Krige &
Krige 1943: 236). Where cultivation of the steep hillslopes has continued, stone
contour walling is still built.
Unworked stones were and still are used in a number of ways for grinding,
crushing, pounding, or smoothing. In a pair of grinding-stones, the lower stone,
lwala, is usually flat, large and fairly rough while the upper stone is smaller,
harder and rounded. Stones are used for grinding sorghum and millet but not for
maize, which is stamped in a mortar. Heads of maize are decobbed on a rough
stone (Fig. 86). During the preparation of clay it is pounded with a wooden
pestle on a large grinding-stone. Smaller, hard stones are used for cracking open
the hard shells of marula pits and generally for cracking nuts. A hard, smooth
LOBEDU MATERIAL CULTURE 145
Fig. 86. Rough stone for removing maize from the cob. Molototsi valley, 1975.
stone, thidél6, is used to compact the surface of a newly made mud floor and
smaller rounded river stones are the traditional smoothers and burnishing-tools
for pottery (see p. 56).
Sandstone, /ehut6, often referred to as the ‘Vim’ stone, is widely used as an
abrasive cleaning agent for utensils. A large grinding-stone on which the lehuté
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
is crushed is generally found near the washing-up area (Fig. 87). A less abrasive
stone is used for scrubbing the body when washing.
Herd-boys in the fields used hollow stones as troughs for goat’s milk which
they curdled by adding acidic fruit juice and ate with spoons made from grass or
leaves (Krige & Krige 1943: 107).
Fig. 87. Grinding-stones for crushing sandstone into an abrasive powder for scouring
utensils. Note pot for water, calabash utensils, and spoon. Modjadji’s village, 1975.
Stone-carving seems to have been confined to the making of soapstone
(steatite) pipe-bowls, gagana (Fig. 88), for smoking dagga, badje (Cannabis sa-
tiva). No technical information was obtained on their manufacture and they are
no longer made. According to Krige (1982: 41) they were used together with a
reed, khetoméla, and a horn water-vessel, naga la khomo. The dagga was placed
in the stone bowl fitted over one end of the reed, the other end was inserted in
the horn filled with water. A glowing ember was placed in the bowl on top of the
dagga and the smoker drew the smoke through the water by inhaling deeply
over the mouth of the horn. This provoked a strong flow of tears and saliva
which could be expelled through another reed, mphasi. This was called maada a
khoho.
By the 1930s dagga-smoking was illegal and the horn water-pipe was no
longer in use. To avoid being arrested people had taken to placing the dagga in
a dampened hole in the ground, and smoking in a lying position. The smoker
filled his mouth with water and drew the smoke from the dagga through a reed,
LOBEDU MATERIAL CULTURE 147
Fig. 88. Stone pipe-bowls, K68, Klapwijk Collection.
khetoméla, positioned to make contact with the dagga which had been placed in
the hole and lit. Nowadays dagga is not smoked through water but in an ordi-
nary pipe or by rolling it in paper.
SUMMARY AND COMMENT
The indigenous Lowveld environment offered a wide range of materials for
possible exploitation. In the above section the uses of clay, grass and other plant
fibres, wood, and stone are described. Although many of the techniques are
termed ‘traditional’ this is not intended to imply a fixed body of skills but to
refer to processes that had developed and continued to develop in a particular
environment and within a self-sufficient subsistence economy.
The form of the material culture at any one time depends on an interaction
of technical, economic, social, and ideological factors, each of which contains a
number of related variables. Technical variables include the types of material,
the tools and the skills; economic variables include the demand for the products,
the division of labour, and the impact of money and mercantile goods on local
production; social variables encompass the range of social contexts for the use of
artefacts and customary patterns of use; ideological variables relate to the sys-
tem of values that gives meaning to certain materials and products.
Natural resources constituted the most important material base in the Lo-
bedu economy. In addition to providing wild foods and land for cultivation and
grazing, the natural environment provided most of the materials on which the
productive technology depended for the construction of settlements and the
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
satisfaction of most other material requirements. Using the Krige Collection as a
sample, over 60 per cent of the inventory of artefacts derive primarily from the
indigenous environment and, if huts and granaries were included, the figure
would be well over 80 per cent.
Knowledge of the natural environment was a very significant aspect of tech-
nology. In the 1930s Krige & Krige (1943: 45-50) recorded the names of over
500 plants used as food and in the technology, and commented that knowledge
of the vegetation and its application in craftwork amounted to a ‘veritable
science’. Selection of appropriate materials for particular purposes was an essen-
tial part of the technical process. This knowledge was acquired from an early age
in play activities and later through participation in subsistence activities. Herd-
boys, for example, developed a sensitive understanding of their environment
and acquired many technical skills. Since children have been receiving formal
education in school, less time is spent in the veld and less is learnt about the
~ natural environment. School education has also tended to undermine a value
system that stressed order in nature and the ritual control of nature by the chief
through the ancestors.
Lobedu orientation toward nature was expressed not only tangibly in the
material culture but intangibly in the rain and fertility cult, and the seasonal
ritual offering of the first termites to the ‘rain-pots’. In the vuhwera and vyali
initiation rituals there was a strong emphasis on the wearing of costumes from
the appropriate natural materials, often associated with rain and damp places
and believed to have positive ‘cooling’ qualities. In the vuhwera costumes there
was an integration of tangible materials and the intangible symbolic values with
which they were invested. These values were further echoed in the songs of ac-
companying vyali girls which were believed to enhance fertility.
Human use of natural resources modifies the environment which, in fact,
becomes partly man-made. The clearing of land for settlement and cultivation
depletes the natural resources and, in the area under study, the accessibility of
many of the traditionally used plant fibres and woods has been appreciably
reduced. Depletion of timber resources prior to the 1930s and consequent legal
restrictions on the felling of trees encouraged a change in the method of roofing
from closely packed to widely spaced rafters, and of hut-building from pole-and-
daga construction to that of mud-bricks. The latter change in building technol-
ogy, which involved a greater use of mud, was also encouraged by an increased
participation of women in hut-building to compensate for the absence of men
working in towns.
Sources of clay, mainly in river-banks, are distributed throughout the area
and in the 1970s were still used extensively by potters. A number of factors have
contributed toward continuity of the craft of pottery. Important among these is
the fact that women are the potters and main users of the products. The technol-
ogy has been passed on from one generation of women to the next, women have
tended to remain in the rural villages, and the pattern of use has remained
relatively constant. Earthenware is generally preferred for cooking on an
LOBEDU MATERIAL CULTURE 149
open hearth and clay pots are less expensive than cast-iron alternatives.
Although water may be carried in lighter tin or plastic containers, it is always
stored in earthenware, which has the practical advantage of keeping water cool
and fresh.
Since the 1930s when only a few of the Christian community made a ‘busi-
ness of pottery’ (Krige & Krige 1943: 320), the manufacture of pottery for sale
has increased. Pottery not only supplies subsistence needs but is one of the ways
in which village women can earn money without having to seek wage employ-
ment. This factor has led to an increase in the scale of production.
Although the staple diet has remained maize-based, there is a qualitative
nutritional difference between home-stamped maize-meal and the less nutritious
refined product bought in the shops. Most people, however, are dependent on
bought maize as over-population has resulted in many people being landless.
The insufficiency of grain is reflected in the disuse of granaries, a reduction in
the frequency of stamping maize and in the decreased use of related utensils
such as mortars, mats for drying meal and pots for steeping grain prior to stamp-
ing.
Changes in basketry since the 1930s seem to relate mainly to reduced
demand for certain products as a result of the changing subsistence base, an
increase of migrant labour, and related changes of values. An example of this is
the kherodwana basket that has become almost obsolete following a change in
the pattern of marriage as a result of migrancy. Formerly this serving-basket was
symbolic of the willingness of a bride to honour her husband and to serve him
respectfully. As migrancy increased many marriages were contracted in the
absence of the man with the result that there was little need for the basket. The
wooden porridge dish, del6, which fitted inside the basket was also used less
frequently. As demand decreased, so did the motivation to transmit the related
technical skills, with the result that they are unknown to the younger genera-
tion.
The replacement of woven beer-strainers by a large wooden sieve was re-
lated to the brewing of maize-meal beer for sale. Formerly woven strainers were
used when beer was brewed in relatively small quantities, mainly from sorghum,
and was used to fulfil social obligations or to show hospitality. This type of
strainer was not suited to straining large volumes nor was it suitable for beer
made from maize-meal. On the other hand, the fine-meshed, wooden-framed
sieve is well suited to straining finely ground maize-meal and can handle large
volumes. As beer-making for sale increased, the woven beer-strainer fell into
disuse.
As migrancy increased, women took over the making of mothatha baskets
and pate mats, traditionally made by men, and they continue to be made by
women for domestic use, although the mothatha baskets have largely been re-
placed by enamelware.
Working with wood, including the weaving of baskets from wood-slivers,
remains exclusive to men and has tended to become more specialized. Many
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
new tools have been added to the tool-kit but the traditional adzes and gouges,
well adapted to their functions, have not been replaced. Tool marks made by a
lehoér6 gouge are visible on a number of items in the Krige Collection and this
type of tool was still used by woodworkers in the 1970s. In general there is a vis-
ible relationship between the tools, the motor action of the user, and the form of
the product. A change in the use of tools is usually detectable on the artefact
and this is particularly so when modern knives, files, and sandpaper are used.
Traditionally many natural materials were used without modification by for-
mal tools. Stones, selected for their particular shape or texture, were used as
nut-crackers, grinders, and smoothers. The abrasive Jehut6 was and still is
widely used for scouring utensils, and the ‘loofah’ plant, leshokhoté (Luffa
aegyptiaca) provides an effective scourer. Many plant fibres can be used as lash-
ings without any technical preparation and the natural shape of the Bauhinia
seed-pod, thema, is well suited to its use as a pottery tool. Thus natural
- materials continue to be used in a variety of ways and have not been replaced by
purchased alternatives.
Despite the transition to a money economy, in many respects the natural
environment remains an important, if diminishing, source of materials and,
where there is still a demand for traditional products, the related technical skills
remain alive.
USE OF DOMESTIC RESOURCES
The products of cultivation and livestock husbandry form the main content
of this section but fruit shells, although natural resources, are included with
gourds, and horns of game animals with those of cattle, as technically they are
treated in similar ways.
GOURDS
Gourds belong to the family Cucurbitaceae, as do pumpkins and melons.
They are cultivated for their fruits and for their leaves which are made into a rel-
ish. The mature fruits of the indigenous species Lagenaria siceraria (formerly
Lagenaria vulgaria) are widely used for making calabash utensils. The smooth-
skinned gourd, thaga, grows in a variety of shapes that lend themselves well to
the making of ladles, scoops and flasks; the small rough-skinned gourd, leraka,
is used mainly for making rattles. The seeds from gourds of useful shape are
kept for the following season.
Seeds are sown during the summer rains between November and January.
Most women plant gourds among the other crops and leave them in the fields
until they are the required size and hardness. By June the mature gourds are
usually considered ‘strong enough’ to be picked and made into utensils. There is
no specialization in this activity—all the village women know how to prepare
calabashes and make those that are needed for domestic use. Some women are
LOBEDU MATERIAL CULTURE Sul
expert at cutting a neat rim and are asked to do this for others. If a woman has
made too many calabashes for her own use she may exchange them for produce
or sell them.
Gourd-working techniques
A gourd of suitable shape is selected for the product in mind. The method
of manufacture is. essentially the same for all calabash vessels. The process ob-
served during fieldwork in the 1970s did not differ from that described by Krige
for the 1930s. A small hole is cut in the gourd which is then cooked in boiling
water for about an hour. When cool, the hole is enlarged, the soft inner flesh
removed with a spoon (formerly with a seed-pod scraper, thema) and the edge of
™
Fig. 89. Repair techniques for calabash utensils. A-B. With wire. C. With string.
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
the rim cut neatly with a knife. The outer skin is scraped off, -thala, and the
gourd is then scoured, -huta, inside and out with powdered sandstone, lJehuto. It
is left outside to dry and is then ready for use. Constant washing with lehuto re-
moves any rough edges from the rim and gives the calabash a smooth finish. In
this way the quality of a calabash utensil improves with use.
The method of preparation differs slightly if the gourd used is too big to fit
into a pot of boiling water. After the hole has been cut, the gourd is inverted
and left for about a week so that the flesh can rot. Thereafter the inside is
cleaned out, the outer skin scraped off, and the surface scoured.
Rattles are made simply by letting a /eraka gourd dry out thoroughly. It is
not boiled. Holes are branded at opposite ends and the inside is cleaned out
with a twig. Small stones or seeds are put into the cavity and a stick longer than
the gourd is pushed through so that it stops the holes and forms a handle at one
end.
Cracked or broken calabashes may be repaired by making a row of tiny
holes on either side of the crack with a fine awl made from a sharpened nail or
umbrella-spoke. The crack is then sewn together with fibre string or fine wire
(Fig. 89A—C). The stitch may be simple oversewing or a more complicated
stitch such as herring-bone. During the months when gourds for making new ca-
labashes are not available, repairs become a practical necessity.
Fig. 90. Calabash utensils: bowl, khethéba, SAM-—9686; scoop, khefhahé, SAM-9711; and
ladle, kheh6, SAM-9693. (All Krige Collection, 1936-8.)
LOBEDU MATERIAL CULTURE 153
Uses of gourds
Utensils and containers
Gourds are made into a number of domestic utensils including bowls,
ladles, scoops and flasks (Fig. 90). A calabash bowl, khethéba, is in daily use for
the staple maize-meal porridge, vhoswa. A smaller khethébana is used for soft
porridge, khepye, and for serving beer individually to an honoured guest. When
used for beer it is referred to as phafhana.
Ladles, zwihd, of various shapes are in regular use. Khehé kha vyalwa is a
long-handled ladle for serving and drinking beer. The long handle makes it easy
to pass from one person to another. Khehé kha mabudu is a short-handled ladle
for light beer not usually drunk in company. Kheho kha meezi is a large ladle for
transferring water from a storage pot to other containers.
An open scoop, khefhahd, is used for handling dry meal. When making
porridge the meal is added to the water with this utensil.
A large vessel, mokhaba (Fig. 91A), is used for transporting liquids. The
small mouth prevents spillage and it is much lighter and less likely to break than
a clay vessel used for the same purpose. In the 1930s a waisted calabash, khe-
khébo (Fig. 91B), was in use as a portable flask for carrying thin porridge.
Seeds or dried beans were kept in similar calabash vessels. Modern containers
have largely replaced calabashes for these functions.
Small-waisted calabashes fitted with stoppers were used as containers for
herbs, medicines, or snuff (Fig. 92). To some extent these have been replaced
by small tins and bottles but they are still used, especially for medicines.
Musical instruments
The resonators of certain musical bows and xylophones consist of suitably
shaped calabashes. The musical bow, khekhaba, (see p. 124) which derives its
name from the North Sotho word for a gourd, has a single calabash resonator
that is held against the player’s body during performance. A xylophone, bela,
(see p. 129) has a resonator below each wooden bar. They are oval in shape,
graded and positioned so that the largest calabash is the resonator for the
deepest note. Near the closed end of each resonator there is a small hole with
resin on the rim and covered with pieces of spider’s web which vibrate when the
instrument is played and add to its tone (Kirby 1934: 50-56). Xylophone players
Kept the seeds of successful resonator gourds so that they could cultivate gourds
of similar shape.
Rattles, tshélé (Fig. 93A), used in rituals of spirit possession are made from
the rough-skinned gourd, /eraka, in the manner described above (p. 152). Leg
rattles, mathodzi (Fig. 93B), are made from the smooth hard-shelled fruit of the
mothodzi tree (Oncoba spinosa). When the lethodzi fruit has turned a dark
brown on the tree it is picked, a small hole is made at each end and the seeds re-
moved with a stick. Small stones are put inside the dry empty shells and a stick
is passed through the holes to close them. Three or four sticks each fitted with
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
B aes
Fig. 91. A. Calabash flask, mokhaba, SAM-9732. B. Waisted flask, khekh6b6,
SAM-9847. (Both Krige Collection, 1936-8.)
LOBEDU MATERIAL CULTURE 155
Fig. 92. Small calabash vessels for snuff or medicines, SAM-—9918 and 9855.
(Both Krige Collection, 1936-8.)
Fig. 93. Rattles. A. Tshélé SAM-9830. B. Mathodzi SAM-9831. (Both Krige Collection,
1936-8.)
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
four or five fruit shells are held together with thongs, or sisal string, threaded
through holes at the ends of the sticks. The thongs are long enough to be used to
tie the rattles to the wearer’s leg. Sometimes slits are made in the shells to en-
hance their sound. They are worn by mal6b6 dancers during spirit-possession
rituals and at recreational dances.
Playthings
Young boys make humming-tops, mankényana, out of lethodzi fruit shells.
A stick is fitted into a hole at one end of the shell and fixed with beeswax. The
string is wound round the stick and when pulled it causes the top to spin with a
humming sound. According to Krige (notes) this is one of the oldest Lobedu
toys. They were seen occasionally during the author’s fieldwork.
Ritual uses
Melons, maguadzi (Citrullus lanatus), were among the first green foods
each year to be ritually ‘bitten’ and offered to the ‘rain-pots’ (Krige & Krige
1943: 274). A small calabash, phafhana, was considered the most appropriate
vessel in which to offer beer to the ancestors.
In former times a doll with a body made of a calabash was given to a girl
when she reached puberty. This doll was kept by the girl until her marriage
when it was carried in a kherodwana basket by the bridal party. If she were not
a virgin the calabash was pierced. According to Krige & Krige (1943: 113) this
custom had fallen into disuse by the 1930s.
SKINS AND HIDES
Livestock husbandry was less important in the Lobedu subsistence economy
than agriculture and this is reflected in the material culture. Goatskins, however,
were important in the traditional technology providing the main material for
clothing as well as for the bellows used in metal-working. Goats were slaugh-
tered on occasions of ritual or social importance and they were also used for the
payment of debts or fines and in many other economic exchanges (Krige &
Krige 1943: 45). Cattle, on the other hand, were rarely sacrificed and the use of
their hides was less significant in the technology.
Skin-working techniques
Skin-working involved a number of processes: flaying the animal, fleshing,
dressing, cutting, and sewing. Men did most of the work but women helped with
the dressing of the skin (Fig. 94). The following description is based on accounts
by men who no longer prepared skins in the 1970s but remembered the tech-
nology.
A skin was removed from the carcass, pegged out with wooden pegs,
dinédo, and rubbed with an abrasive stone or dry maize cob to scrape away the
LOBEDU MATERIAL CULTURE 157
Fig. 94. Woman applying fat to a goatskin. Photo: E. J. Krige 1936-8.
fat and clean the flesh side (Fig. 95). This removed matter that would putrify
rapidly if the skin were left untreated. Thereafter it could be allowed to dry out
before being softened or dressed. At this stage the skin was still very susceptible
to infestation by insects or bacteria and the experienced skin-worker knew the
value of certain plant oils and repellents that could be used to protect the raw
hide (Krige & Krige 1943: 47). The dressing-process that followed rendered the
skin less prone to decay but did not make it completely stable under unfavour-
able conditions.
The pelt was dampened with warm water for the softening-process, -suha,
which was done by rubbing the skin briskly by hand. This was an arduous pro-
cess and a number of men would work together on a single skin or take turns in
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 95. Goatskin drying after the flesh side had been cleaned, Molototsi valley, 1976.
working on the skin. Pips of the ditshidi fruit (Ximenia caffra) were ground be-
tween two stones to extract the fat, which was then applied to the skin. This
acted as a dressing and kept the skin supple. The hair was usually left on
goatskins but could be removed before softening by soaking the skin in water
and scraping the skin with a rough stone. Cattle-hides used for sandals, shields,
and quivers were not softened but covered with dung and buried so that the hair
would rot and fall off.
For certain purposes a prepared skin had to be cut and sewn to a conven-
tional pattern. This was the work of a specialist craftsman. A woman’s skirt,
motshe (Fig. 96), required particular skill. Gussets were cut to give the skirt the
correct shape; flaps of skin, dinyedi, were cut and sewn on later for decoration;
a triangular ‘tail’ was cut and sewn on to the waist so that the hairy side faced
outward when folded over the back of the skirt. Pieces of skin were formed into
small studs and attached so that the hair showed up against the flesh side on
which a nap was raised by making small incisions with a special metal tool,
mosédo (see p. 177), with a cutting edge that could be sharpened. The nap-
raising process, -déga, gave the skirt its characteristic texture and without this
finish a skirt was not considered complete. Sewing the pieces together was done
by passing sinew through holes made with a small awl. Ties at the waist of the
skirt were traditionally made of plaited wild-cotton thread (/etshida) and later of
cotton thread bought from traders.
LOBEDU MATERIAL CULTURE 159
\ VAN ‘
t\ \ yi
\ \ \B
\s DAN na ayy
Vad SAV yet
i. \\
oe
Fig. 96. Skirt, motshe, for a married woman, SAM-9762, Krige Collection, 1936-8.
A number of factors led to a decrease in demand for skin products and a
decline in skin-working technology. The early adoption of European dress by
migrants, the influence of the mission, and the popularity of cloth meant that
fewer people wished to wear skin clothing and fewer men learnt the traditional
skills. Many young men left the rural villages to work in town and the money
they brought back to the villages had a compounding effect on the decline of
skin-working by replacing goats in many traditional transactions and thus reduc-
ing the supply of skins (J. D. Krige 1934: 137, 145; Krige 1941: 17).
Uses of skins and hides
Clothing
Traditionally men wore a loin-covering, thséte, which was a triangular piece
of goatskin that passed between the legs, and strings from the three ends were
fastened in a knot at the back of the waist. A second piece of skin, called
bokone, was attached to the waist-string to cover the buttocks. No other cloth-
ing was worn except a goatskin cloak in winter. In the 1930s it was very rare for
men to wear skin clothing although it was not uncommon for a skin loin-
covering to be worn under a pair of trousers (Krige 1982: 29).
Women’s clothing consisted of a small triangular front apron, thethd, and a
back apron, thebyana (worn mainly by unmarried girls), or a skirt, motshe, worn
by married women and by unmarried girls (Fig. 97). A girl initiate coming out of
seclusion at puberty was given a front apron, a back apron, and a skin cape to
wear over the shoulders. A photograph taken in the 1930s (Krige & Krige 1943,
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 97. Young girls wearing skin skirts and waist ornaments. Also note ear ornaments and
hair-styles. Photo: E. J. Krige, 1936-8.
pl. 8a) shows that the hairy side of the skin was worn toward the body and the
other side has a raised nap similar to that found on the motshe.
Skin aprons were important in the traditional exchanges that preceded a
marriage. Among the gifts from the boy’s to the girl’s family was a goat, pudi ya
kebolao, that provided the skin for the groom to prepare a thebyana for his fu-
ture wife. This and a thethd, also prepared by the groom, was worn until after
the birth of her first child. Thereafter the thebyana was replaced by a motshe.
These skin aprons were more than simple items of clothing, they expressed ma-
terially the relationship between the two families. Thus ‘People not only create
their material culture and attach themselves to it, but also build up their re-
lationships through it and see them in terms of it’ (Evans-Pritchard 1940: 89).
By the 1930s cash payments had replaced many of the traditional gifts
(J. D. Krige 1934: 137). Payments of money were explained in terms of ‘thank-
ing’ the girl’s parents for consenting to the marriage. In many cases the groom
was working in town and absent from his village at the time when the prelimi-
nary arrangements for his marriage were being undertaken.
The production of skin clothing steadily declined. Imported cotton clothe
such as salempore and morina had been introduced by traders in the area from
the mid-nineteenth century onward and, together with mission influence and
LOBEDU MATERIAL CULTURE 161
changing economic conditions which encouraged migrancy, led to the eventual
replacement of skin clothing by cloth wraps for women, and trousers and shirts
for men. In the 1970s salempore and morina cloths were still worn by conserva-
tive women. Small skin aprons were retained by old women as under-garments
but most young women wore modern dresses in keeping with their modern life-
style. Skin clothing for general use was something of the past but the tradition
was preserved to some extent in initiation schools. Skin skirts were also worn as
special costume for dances or celebrations.
Hide sandals, zwiéta zwamphashane, were rare in the 1930s and today they
are not made at all. A piece of hard hide was cut to the size of the foot and
strips of skin were passed through slits in the sole at the toe and at the heel to
fasten the sandal to the foot.
Sandals made of rubber from motor-car tyres were popular in the 1930s and
were said to be durable and more comfortable than hide sandals. In the 1970s
these sandals were still made and worn fairly widely.
Baby-slings
A skin sling, thari, was used for carrying a baby on the back (Fig. 98). Tra-
ditionally the skin used for making a thari came from the goat that a husband
slaughtered in honour of the new-born baby. The skin was dressed in the man-
ner described above and no cutting was needed. A nap was raised on the flesh
side using the metal tool, mosédd. The hairy side was worn on the inside against
the body and a flap was folded over on to the side with the raised nap. To this
flap various things might be attached. A thari collected in the 1930s has attached
to it a soft fibrous bulb, /e6me, which was used to wipe the baby’s nose, a small
skin bag of protective medicines, and strings of coloured glass beads, which ex-
pressed the mother’s joy in her child. Most of the skin baby-slings seen in use in
the 1970s had been handed down from previous generations but they were oc-
casionally still made when a child was born. Most women, however, used blan-
kets for carrying their babies.
Bags and pouches
Bags for general use and for special purposes were made of skin. A reptile-
skin pouch, khesigo, was used to keep divining-bones when not in use. Medi-
cines were often carried in small skin bags, dithévhélé, which could be attached
to a girdle or baby-sling. Large skin bags used by men served the purpose of
pockets before trousers were widely adopted (Krige 1982: 30). A set of reed
pipes was Kept in a skin bag made specially for this purpose.
Traditionally snuff was kept rolled up in a piece of soft goatskin, moséha
(Fig. 99), or in a calabash snuff-container (see p. 153) carried in a small pouch
made of dassie skin. Horn snuff-boxes (see p. 167) were introduced by Tsonga-
speakers and in the 1930s were widely used by Lobedu men (Krige notes). Small
tins and bottles were and still are used as snuff containers. In the 1960s a piece
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 98. Young girl carrying a baby in a thari. Photo: E. J. Krige,
1936-8.
Fig. 99. Roll of skin for holding snuff, moséha, SAM-9854, Krige Collection, 1936-8.
LOBEDU MATERIAL CULTURE 163
of plastic sheeting rolled up in the manner of the moséha was a popular way of
carrying snuff (Krige 1982 pers. comm.).
Poisoned arrow heads were kept in a hide quiver, khéd6b6, and sharp
knives were kept in a hide sheath, khelada.
Shields
An ox-hide shield, khetagu, was described by an old Lobedu man as being
roughly of hour-glass shape with the narrow section at the grip. No specific in-
formation on making shields was obtained and no examples were seen in the
field or in museum collections.
Skin components of weapons
A sleeve of tail skin from an ox or cow was used for the attachment of a
spear-head to the shaft (see Fig. 112C). The wet skin was pulled over the join
between head and shaft and when dry it formed a tight band that secured the at-
tachment.
The string of a bow was made of a strip of twisted hide (see p. 115).
Thongs
Thongs or riems, mazwa, were used mainly for tethering animals and for
inspanning cattle, a practice adopted from white farmers. They were also used
generally for purposes that needed a strong binding-material. They are still
made when suitable hide is available.
To make a riem, an ox-hide is put into water to soak and when thoroughly
wet is put flat on the ground and cut round and round to give a very long strip of
hide. The strip is thrown over a strong branch or a frame and a heavy stone is
tied to the ends to weight them. In the 1930s bored stones were used for this
purpose (Krige 1982: 31). Using a long stick, the thong is wound up tightly and
then allowed to unwind. This is repeated a number of times while the skin dries
out. The friction causes most of the hair to fall off and fat is applied to help pre-
serve the skin and prevent the thong from becoming brittle.
This method of making thongs is widespread throughout southern Africa.
Other uses of skins and hides
Bellows
Goatskin bellows were used for forcing air into the furnace or forge during
the smelting and smithing process. The two ‘bags’ of the bellows were made
from the uncut skin of the hind legs of a goat (Krige 1982: 31).
Drums
Cow-hide was used for the heads of géma and thithimedzhé drums and
goatskin for the tambourine, khethadada (see p. 123).
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mats
Unworked but cleaned goatskins were used by some as sitting-mats. They
were not as common as grass mats (Krige 1982: 31).
HORN AND BONE
The care of livestock was predominantly a male activity as was work on
horns and hides. No specialized skill was required to convert horns into con-
tainers of various kinds and most horn artefacts utilize the natural form of the
horn without any additional working. No artefacts were carved entirely from
solid horn. Horn snuff-boxes were made by using both the solid end-section and
the natural hollow of the horn.
Bone was not used to a large extent and when it was used the natural shape
of the bone, or piece of bone, was modified to suit the particular function. Solid
bone was not used as a carving medium.
Uses of horn and bone
Medical equipment
1. Cupping-horn
A cupping-horn, mothoho (Fig. 100A), consists of a small horn or end-
section of a larger horn perforated at the solid tip so that it is open at both ends.
Beeswax (modu) is used to seal the narrow opening during the cupping-process.
Cupping or blood-letting, -loméha, is practised to cure headaches or pains
believed to be caused by impure blood. Most people know how to use a cup-
ping-horn and the operation does not have to be carried out by a diviner. Two
small incisions are made in the affected area over which the wide end of the
horn is placed. Air is sucked through the narrow end and blood is drawn into
the horn by the vacuum. The tip is sealed with wax and blood continues to flow
into the horn until the operator breaks the vacuum and removes the horn. The
blood is buried in a secret place for fear of it being used in witchcraft.
2. Medicine-horns
Herbalists store medicines in ox-horns (Fig. 100B). Rain-water for use in
special rituals is also kept in horn vessels. Horns used in this manner are not
carved but many have holes drilled near the rim for the attachment of a thong.
3. Divining-bones
A set of Lobedu divining-bones consists of about forty pieces, the most im-
portant of which are the four flat carved bone or ivory pieces, two male and two
female, which fall in sixteen different combinations, the cardinal positions. The
set includes a number of pieces not made of bone, for example, shells, river
pebbles and marula pits, as well as pairs (male and female) of bones from ani-
mals associated with totems. The basic principles of divination are outlined by
Krige & Krige (1943: 226-227).
165
LOBEDU MATERIAL CULTURE
ine-horns at
1c
. Med
B
llage, 1973
horn in use, Headman Mohale’s vi
Headman Mohale’s village, 1973.
Fig. 100. A. Cupping
S Vl
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
Musical instruments
1. Trumpet
The traditional horn for making a phalafhala trumpet (Fig. 101A) was that
of the sable-antelope from which the instrument derives its name, but kudu
horns were also used. The horn was left to dry out until the core became loose
and was easy to remove. An embouchure was carved in the side of the horn at
the point where the hollow is narrowest, i.e. just before the solid tip starts. If
the horn showed signs of splitting it was strengthened with a skin sleeve to pre-
vent further cracking. Horn trumpets are seldom made today but a number of
old ones, belonging to district headmen, are still in use.
The phalafhala produces only one note but when blown hard it is loud and
carrying. It was and still is associated with authority and it sounded as a signal or
to make important announcements, for example to warn of a veld fire, to sum-
-mon people to meet at the capital, to herald the arrival of a party of gosha
dancers, or to announce the arrival of tribute sent to the Queen.
2. Whistles
Small antelope horns were made into whistles, dinaga (Fig. 101B). The
solid core was removed, leaving the horn-sheath closed naturally at one end and
open at the other. This formed a simple stopped pipe and sound was produced
by blowing across the open end. Horn whistles were used in hunting and by
herd-boys. They are seldom made today as game is protected and boys spend
relatively little time herding or playing in the veld. Whistles were also made
from bone, often the hollow tibia of a small animal. One end was stopped and
the whistle was played by blowing across the other end. According to Krige &
Krige (1943: 135) during the vyali the Bird of Muhale communicates by
whistling through an instrument made of bone but the Bird is also said to use the
mphenyi made of reed (see p. 99).
Fig. 101. A. Horn trumpet, phalafhala SAM-9822. B. Horn whistle, nagana, SAM-9818.
(Both Krige Collection, 1936-8.)
LOBEDU MATERIAL CULTURE 167
Pipes
In the past dagga, badje, was smoked through a water-pipe that consisted of
a stone pipe-bowl, gagana (see p. 146), a reed, khetoméla, and an ox-horn
water-vessel, naga la khomo. The ox-horn did not require special preparation.
The dagga was placed in the bowl and the reed formed the pipe-stem that was
inserted into the horn of water so that the smoker could draw the smoke
through the water when he inhaled deeply at the mouth of the horn.
Snuff-boxes
Horn snuff-boxes were introduced by Tsonga-speakers and their use was
adopted mainly by Lobedu men. The form of the snuff-boxes (Fig. 102A-—B) fol-
lows the natural shape of a horn. A hole is bored from the existing hollow
through the solid tip of the horn. The wide part of the opening is then closed
with a plug of wood or gourd and the narrow mouth is fitted with a small stop-
per. It is not uncommon for a horn snuff-box to have a carved ridge near the
mouth and a thong tied below the ridge. In an unusual example (Fig. 102A) col-
lected in the 1930s a human figure has been carved following the natural shape
of the horn.
Tools
1. Rolling-tool
An ox-horn, flattened on one side, is used as a tool, khehdthod, to coil wire
round a foundation of tail hair in making wire arm- and legrings, maséga.
Fig. 102. Horn snuff-boxes.
A. SAM-9850. B. SAM-—9851.
(Both Krige Collection, 1936-8.)
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
The tail hairs, maditsi, are first cut, twisted, and the ends joined to form a
circle. The wire is passed through a small split stick and wound a few times
round the hair circlet, which is then placed on a wooden plank and stroked,
-hota, in a downward movement with the horn tool so that the wire rolls evenly
on to the base of hair (Fig. 103). In the 1930s brass or copper wire was bought
from traders and today this had been replaced by aluminium wire. Before wire
could be bought it was made by an arduous process of drawing hot metal
through a perforated plate (see p. 172).
Traditionally women wore large numbers of maséga on their ankles (see
Fig. 109) and wrists but these ornaments have become less common. Legrings
are said to decrease mobility and there is an increasing tendency for young
women to adopt European dress.
Fig. 103. Horn tool used to roll wire on to foundation of tail hair in making maséga bangles,
Molototsi valley, 1976.
2. Marula tool
A bone tool, /efhéd6 (Fig. 104), is used to remove the tough skins of the
‘marula fruits from the flesh prior to making marula-cider, mokhobe. The exist-
ence of a special tool for this purpose and of a special bowl (see p. 64) for
drinking marula-cider is an indication of the importance of the marula fruit to
the Lobedu. The fruit ripen in late January and February when crops are imma-
ture and there is little grain left over from the previous harvest. The marula
season is, therefore, eagerly awaited. Marula-cider is rich in vitamin C and a
source of nourishment as well as being a refreshing drink. It is usually made by
women in the fields under the marula trees where it is also consumed with much
enjoyment.
LOBEDU MATERIAL CULTURE 169
Fig. 104. Bone tool, lefhéd6, for removing the skin from marula fruit,
SAM-9746, Krige Collection, 1936-8.
The ripe fruits that have fallen to the ground are used to make the cider.
The tough skins are removed with the /efhéd6 and the juice is squeezed from
the flesh with a kneading action, -soha. The pulp and the pits are placed in a
large clay pot, thukhwana. Water is added and the fruits are rubbed, -suha
marula, to extract all the juice, which is then ladled into another pot, leaving the
pits behind.
At this stage the juice, lesebeta, is left to ferment. After one day it is still
very sweet and is known as /edutu. A layer of scum, ledobe, which has risen to
the top, is removed. A bowl is placed over the mouth of the pot and the edges
are sealed with some of the /edobe. Another layer of scum rises to the top of the
liquid and is removed before the cider is ready for drinking. The longer it is al-
lowed to ferment, the less sweet and more intoxicating the cider becomes. It is
usually allowed to ferment for about three days before it is drunk.
Water is added to the pits that remained behind and is decanted on the fol-
lowing day to provide a dilute children’s drink, malithele. Thereafter the pits are
left to dry and are cracked open later for the kernels, which are ground and
added to vegetable relish.
3. Plastering-tool
The flat rib bones of cattle are recorded as having been used at Medingen as
trowels when plastering walls (Klapwijk Collection, Transvaal Provincial Mu-
seum Service K108A).
DUNG
Cattle-dung was mixed with mud to form a kind of cement used in hut con-
struction for the plastering of walls and floors and for hardening the floor of the
courtyard. The mixture of dung and mud was spread over the floor surface and
compacted with a wooden beater (see Fig. 55) or a smooth stone, thidéld. In the
1930s decoration of floors and walls was rare (Krige 1978 pers. comm.) but in
the 1970s finger-patterns made while the last layer of dung was still wet were
very popular at the capital and elsewhere in the capital district, and gave an at-
tractive finish to the floor.
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
Formerly dung was used to line the inside of grain pits (see p. 140) and to
fill in the spaces between the reeds of the door of a cooking-hut (see p. 85).
Dung was not generally used as fuel nor was it widely used as a fertilizer.
Although it was realized that adding dung to the soil improved the crop yield, it
was said also to encourage the growth of weeds, which discouraged its use
(Muneri Modjadji to Holloway Commission 1930).
SUMMARY AND COMMENT
Domestic resources, the products of cultivation and livestock husbandry,
are secondary resources in that they depend on the natural resources in the form
of arable land and grazing, and on agricultural and pastoral technology. In the
above section the uses of gourds, skins, horn, and bone are described. Fruit-
shells and horns of game animals, although natural resources, are included as
their technical preparation is similar to that of gourds and cattle-horns respect-
ively.
In general the processes for which women were traditionally responsible
have remained more stable than those carried out by men, and this is so in the
case of calabash-making. There is no specialization in this craft as most women
know how to cultivate gourds and make a variety of utensils. The continued use
of calabash vessels for serving porridge seems to be related, as in the case of
cooking-utensils, to the constant dietary pattern and the continued demand for
calabash utensils well-suited to their varied domestic uses. It is not uncommon
for enamelware to be used concurrently with calabashes for storing dry foods
but calabash vessels have not been replaced for serving porridge, decanting
water and, in some situations, for drinking beer.
The long-handled beer-ladle, which is passed from person to person at
social beer-drinks, can be seen as expressing generosity and good social rela-
tions. At beer-drinks where bought beer tends to be consumed by individuals,
the calabash ladle is not appropriate and the glass jars or tins of standard
volume that are used instead reflect materially the commercial principle under-
lying this kind of beer-drinking.
. The most important product of livestock husbandry that was used in the tra-
ditional technology was goatskin. This provided the basic raw material for skin
clothing. Men processed the skins and manufactured the garments for domestic
use. By the early twentieth century, however, production of skin clothing had
started to decline. Migrancy imposed the need for men to adopt European
dress, the mission influenced the clothing of those who adopted the faith, and
these early pressures coincided in the late 1890s with the rinderpest, which
almost annihilated all the livestock of the area.
Migrant labour had a further effect on the decline of skin-working by reduc-
ing the number of craftsmen. Money that migrants brought back into the area
had a compounding effect by replacing goats in many traditional transactions,
thus reducing the supply of skins.
Well into the twentieth century women continued to wear skin clothing as
LOBEDU MATERIAL CULTURE 7A
well as the cotton cloth wraps that had become ‘neo-traditional’ since their wide-
spread adoption and popularity in the nineteenth century. Even as the use of
cotton clothing increased, older women often wore a skin apron underneath a
cloth skirt or wrap. By the 1970s, however, skin clothing had fallen almost en-
tirely into disuse. Old skin skirts were brought out for ceremonial occasions but
new ones were seldom made and few men remembered the former techniques of
making skin garments.
Among the changes that paralleled the move from skin to cotton-cloth
clothing were the replacement of the goatskin baby-sling by a lighter cloth ver-
sion that did not have the disadvantage of being smeared with fat, which was
absorbed by the wearer’s cloth wraps, and the decline in use of skin bags by men
for carrying personal possessions in favour of trouser pockets.
Horns traditionally used as containers or flasks, especially for medicines,
were still used for this purpose in the 1970s. The rain medicine, mofugo, used in
a ritual of appeasement in 1973 (see p. 101) was carried to the site of the cer-
emony in an ox-horn flask and returned afterwards to the home of Modiga who
was responsible for its safe-keeping.
Horn also continues to be used in the form of the tool khehdéthé, made
from a flattened horn-core and used for coiling wire round a circlet of hair to
make a bangle. This tool is so well adapted to its function that it is difficult to
think of an alternative that would work as well. Equally well adapted is the bone
tool, /efhéd6, used to remove the tough outer skin of marula fruit prior to mak-
ing mokhobe cider. The use of a specific tool for this purpose and of a special
bowl, kheritswana (see Fig. 13C) for drinking mokhobe is an expression of the
social and dietary importance of the marula fruit.
Although not as extensively exploited in the technology as the natural
environmental resources, domestic resources supplemented the indigenous
materials and were essential to the traditional material culture.
USE OF IMPORTED MATERIALS
METAL
Metal ores were not mined in the Lobedu area. The closest source of iron
ore, gwedzi, is some distance to the north-east, and when metal-working was an
active industry ore had to be carried on foot to the Lobedu smelting-sites. Krige
(1941: 9) recorded that unfortunates who had no other way of earning hoes or
cattle to acquire a wife carried iron-ore on foot-paths from a ‘long way to the
north’ (probably the Tshimbupfe area) in return for a hoe per journey. The
location of smelting-sites some distance from the ore source is not unusual in the
Transvaal Lowveld. Archaeological evidence from the Phalaborwa area indi-
cates that ‘it is clear that smelting-sites were located near the centres of demand
and distribution, rather than near the ore-supply’ (Van der Merwe & Scully
L971 182).
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
Copper came into the Lobedu area from Messina and Phalaborwa, which
were important centres of copper production in the pre-colonial period (Wilson
1969: 173-174). According to Van Warmelo (1940: 4) until the mid-nineteenth
century there was ‘a constant stream of travellers going to and coming from the
copper mines at Messina, where they bartered for the precious metal’. Tin is
known to have been mined in the Rooiberg area and may have been traded over
a wide area.
Early trade through Delagoa Bay was a source of iron, copper, and brass,
and in the late nineteenth and early twentieth centuries European trade goods,
including iron hoes, tools, and wire, were brought into the area in much larger
quantities than before. During this period and later into the twentieth century
another source of iron and metal goods was importation by migrant labourers
returning from work on the mines.
_ Metal-working techniques
‘In the old days . . . iron making was one of the great industries’ (Krige &
Krige 1943: 47). By the 1930s this industry had ceased largely because of two re-
lated factors—the appropriation of the ore sources by whites and the introduc-
tion of cheap factory-made metal objects, often imitations of the indigenous
products, which rapidly undermined local production.
A number of Lobedu smelting-sites are said to exist. So far none has been
excavated but pieces of iron-ore, gwedzi, and slag from the furnaces were col-
lected by Krige (SAM-—9889, 9890) from a terraced site near Modjadji’s Nek. At
this particular site there is also clear evidence of habitation in the form of hut-
floors and daga. No historical record of Lobedu smelting-technology has been
found by the author and men interviewed during fieldwork had only the vaguest
memory of the processes involved. It was said that the siting of a furnace was
important both in relation to the fuel supply and the prevailing winds. It was
also necessary to know the types of wood that would produce very hot coals.
Goatskin bellows with clay nozzles were said to have been used to keep the fire
burning for many hours but technical details were not remembered.
Metal-working was a specialized industry and there is no record of the ex-
tent of Lobedu involvement in the production of metal goods. The Lobedu ob-
tained many of their metal artefacts through trade with the Phalaborwa, Venda
and Lemba, and locally from Lemba craftsmen living in the Lobedu area. The
tools (Fig. 105A—D) of a Lemba smith who was over 80 years old in the 1930s
are preserved in the Krige Collection. They include a hammer, nddo, used for
forging metal implements; tongs, Jomand, for handling the red-hot metal; a
draw-plate, Jeg6khd, used in wire-drawing; and a vice, bagd. Also in possession
of the smith was a tin ingot, /ekatea, which was said by him to have been used in
‘joining metals’.
Wire was made by heating a piece of soft metal, usually copper or bronze,
and shaping it so that it could pass through a hole in the draw-plate. The vice
securing the piece of metal was wedged in the fork of a tree while the wire was
LOBEDU MATERIAL CULTURE 16/3
D
Fig. 105. Tools that belonged to a Lemba smith. A. Vice, bag6, SAM-—9898. B. Draw-
plate with holes of different size for wire-making, legd6khé6, SAM-9896. C. Hammer,
nod6, SAM-9895. D. Tongs, lomanéd, SAM—9894. (All Krige Collection, 1936-8.)
drawn out with the plate. According to Stayt (1931: 65) a forked stick was used
to lever the plate as the wire was drawn out and wound round the tree. For thin-
ner wire the process was repeated using a smaller hole in the plate.
Uses of metal
Hoes
Before the introduction of the plough, hoes, mazhébé, were the most
important agricultural implements and their use has continued in areas unsuit-
able for ploughing. In addition, hoes were important as exchange media especially
in marriage contracts. Hoes so used established legally binding obligations
between the kin groups involved (Krige 1964: 162).
Hoes are said to have been forged locally but many were traded from neigh-
bouring people. In the early nineteenth century the Venda were exporting hoes
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 106. Woman hoeing in the fields. Photo: E. J. Krige, 1936-8.
‘in thousands to neighbouring districts’ (Beuster 1879: 239) and the Phalaborwa
people were also trading extensively in hoes. It is possible that specialists con-
trolled the production of hoes as in the case of a Phalaborwa iron-worker who
‘taught others to produce iron but retained exclusive monopoly over the knowl-
edge of hoe-making’ (Scully 1978: 254). Although there must have been fluctua-
tions in the exchange rate of hoes through time and in different social and |
economic circumstances, it was only after the large-scale introduction of cheap |
factory-made hoes in the late nineteenth century that the traditional value of
hoes seriously declined.
The form of hoe-heads collected in the Lobedu area in the 1930s is roughly
diamond-shaped with a faint midrib running down the centre. Venda hoes illus-
LOBEDU MATERIAL CULTURE 7S
trated by Stayt (1931, pl. 29) are of similar shape. For use in the fields (Fig. 106)
hoe-heads were attached to a long wooden handle. In general it seems that hoes
used in agriculture did not differ in shape from hoes used for exchange, except
that the latter were new and not worn down through use. There is some evi-
dence, however, of heavier, elongated spade-shaped hoes being used solely for
exchange. “These hoes were too large and heavy for tilling the soil and were
merely kept in a hut’ (Krige 1964: 162).
By the 1930s hoe-heads (Fig. 107) of local manufacture had come to be re-
garded as heirlooms that had links with the ancestral spirits. They were often
partly embedded in small clay-mound shrines where offerings were made to the
ancestors.
Fig. 107. Old hoe-head kept for use in ancestor
rituals, Mohale’s village, 1973.
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Ornaments
Copper and its alloys were used for making solid neckrings and bangles,
mefhiri (Fig. 108A), beads, zwirogaana (Fig. 108B) and coiled wire arm- and
legrings, masega (Fig. 109). Ornaments made of hand-forged metal were rare
even in the 1930s and are now used as amulets and regarded as ‘thugula’ objects.
Kherogaana beads of the ‘wrap-around’ type, i.e. made from a narrow strip of
hand-worked copper cut and bent to form a circle, are among the oldest Lobedu
ornaments. Individual metal beads in the Krige Collection (SAM-—9910) are said
to have been handed down from generation to generation as highly valued
objects. Similar beads, dithotho (SAM-—9911), made in the same way from factory-
produced copper and brass were much more plentiful and less valued than the
ancient copper beads. Dithotho beads were strung in large numbers on cotton
thread to form beaded strands with tassels at the ends and were worn attached
Fig. 108. A. Solid metal bangle and neckring, SAM—9908 and 9909. B. Wrap-around metal
beads on thread foundation, SAM-9913. (All Krige Collection, 1936-8.)
LOBEDU MATERIAL CULTURE M77
Fig. 109. Rolled wire legrings, maséga, Modjadji’s village, 1973.
to skin skirts. More rarely the older kherogaana beads were worn in strings by
those who had enough of them (Krige 1983 pers. comm.).
The mefhiri neckrings and bangles collected in the 1930s are made of brass,
which suggests that they are not as old as the copper kherogaana beads although
they are said to have been of local manufacture. They were rare in the 1930s and
in the 1970s were not seen in use at all.
Coiled arm- and legrings are still made (see p. 167) but wire of copper and
brass has been replaced by aluminium wire and these ornaments are not worn in
such large numbers as in the past.
Tools and weapons
When metal-working was an active industry the iron heads for tools and
weapons were locally produced. Later scrap metal was used and forged into the
required artefacts. More recently, purchased tools and weapons have come into
use. Many of the traditional tools used in wood-working (see p. 101) and skin-
working (see p. 158) had characteristically shaped iron heads that were related
to their particular technical uses. The action of the craftsman in handling the
tool was taken into account when forming the head and when hafting it. The
long-handled gouge, /ehdr6 (Fig. 46), is a good example of this.
The following iron-headed tools and weapons (Figs 110-112, see also Fig.
61) were collected in the 1930s: adze, bédwana; gouges, mahdér6; poker, moro;
awl, morogola; nap-raiser, mosédo; knife, mofhaga; spatula, lehobd, for wiping
perspiration from the face; spear, lerumd; axe, zaga; and arrow-heads,
mesévhé. Artefacts, locally smelted and forged, were rare in the 1930s. They
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 110. A. Poker, mor6, SAM-9725. B. Awl, morogola, SAM-9742.
(Both Krige Collection, 1936-8.)
ue
Aut
Fig. 111. A. Knife, mofhaga, SAM-9848. B. Sweat-scraper, leh6b6, SAM-9887.
C. Skin-working tool, moséd6, SAM-9764. (All Krige Collection, 1936-8.)
were highly valued as thugula heirloom objects even if rusted and in bad repair
and they were thought of as links with the ancestors. These beliefs are still held
by many Lobedu.
GLASS AND SHELL
A distinction should be made between Lobedu ancestral beads, which prob-
ably pre-date the eighteenth century, and glass beads of more recent origin.
Ancestral beads are said to have been brought from north of the Limpopo River by
the founding ancestors, an oral tradition which is substantiated by comparative
chemical analysis of similar beads excavated at the Rozwi ruin at Dhlo-Dhlo
(Zimbabwe) which was occupied around ap 1700 (C. Davison & Desmond
Clark 1976: 132-137). Lobedu ancestral beads include those known as modala,
modzike and thanganwe, that are similar in appearance and in chemical
LOBEDU MATERIAL CULTURE 179
Abe?
Ret Se
EG Wa
“Qs
ee
eG
= "Ge .
Fig. 112. A-B. Axe-heads, SAM-—9884 and 9885.
C. Spear-head and butt, SAM—9843. Note skin sleeve
for attachment of head to shaft. (All Krige Collection,
1936-8.)
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
composition to the Venda ‘beads of the water’, vhulungu vha madi, described by
Van Riet Lowe (1937: 368), Dicke (1937: 406), and Schofield (1958: 189-192).
Ancestral beads, presented by Krige in 1937 to the Archaeology Department of
the University of the Witwatersrand, were seen by the author. They include
blue-green, modala, cobalt blue, modzike and thanganwe, red, mokhwibidu,
green, lethodi, yellow, thaha, and opaque white, khefhago. Many are small cane
or snap beads having a waxy, translucent quality. The dark cobalt-blue beads
tend to be larger. These beads and other ancient beads were and still are
regarded by the Lobedu as thugula or sacred objects.
Fig. 113. Beaded ornaments. A. Necklets, khekhadwa, SAM-9781, and leda-
bara la dithséhéla, SAM-9780. B. Armbands, zwifhédd6, SAM-9782. (All
Krige Collection, 1936-8.)
LOBEDU MATERIAL CULTURE 181
Glass beads acquired through trade in the late nineteenth and twentieth
centuries differ in appearance from ancestral beads in lacking the distinctive
translucent quality and characteristic colours; they are also far more abundant.
Whereas heirloom beads were usually treasured individually and used as amu-
lets, later beads were very plentiful and were made into a variety of ornaments.
Nineteenth-century cane beads were made into a many-stranded necklet,
khekhadwa (Fig. 113A), which was worn traditionally by a young bride when
she first came to her husband’s home. This necklet was also worn by recently in-
itiated boys and girls in recognition of their new status (Krige 1982: 34). Charac-
teristically the khekhadwa is strung so that it has four bands of colour composed
of many strands of beads, usually green, madalayane, blue, moddgwa, yellow,
khirulwana, and black, lesoli. Also worn traditionally by a young bride (Krige &
Krige 1943, pl. 12b) were zwifhéd6 armbands (Fig. 113B). The beads used were
small, rounded oblates, mainly black, white, and dusky pink. They were strung
in rows between pieces of hide about 13 cm in length. Old examples were strung
on sinew; in more recent specimens cotton string has been used. Zwifhéd6 arm-
bands were also worn for dances and celebrations, together with waist orna-
ments, zwibébédana, which consist of multi-coloured beadwork flaps worn over
a skin skirt (Krige & Krige 1943, pl. 7b). These bead ornaments are no longer
made but many Lobedu women still have old pieces in their possession. The
large number of examples in the Klapwijk Collection suggests that they were
widely used well into the twentieth century.
In the late nineteenth and early twentieth centuries ostrich eggshell beads
were brought by traders into the Lobedu area. Migrants returning from the
Fig. 114. Girdle of ostrich eggshell beads, mofthaga, K102A, Klapwijk Collection.
182 ANNALS OF THE SOUTH AFRICAN MUSEUM
mines could buy these beads in the Munnik and Soekmekaar areas which were
said to have been centres of manufacture (Klapwijk Collection K102A, C) as
were the Phalaborwa and Venda areas. Beads could be bought loose or in
strings, some were roughly cut, others smoothly polished, and they were made
up into mothaga girdles and worn by Lobedu women and young girls. The mo-
thaga (Fig. 114) is a single or double string of ostrich eggshell beads with a single
seed of the wild banana (modolo) at the centre back. Worn over a skin skirt tied
in the front with the seed always at the back, the girdle was thought to have pro-
tective qualities for the wearer and to enhance fertility. They were made in large
numbers and from 1900 onwards factory-made white glass beads were imported
by traders to replace ostrich eggshell beads (Klapwijk Collection K102A).
Cowrie shells from the coast were traded in the Lowveld and worked into
ornaments (Fig. 113A). They were thought to bring good fortune to the wearer.
The cylindrical central spire of the cone shell was made into an ear ornament,
dibadana (Figs 115-116). It was worn by women through the ear lobe in the
manner illustrated by Tyrrell (1968: 62). Early dibadana are said to have been
made of ivory (Krige 1982: 35); the shell version replaced the ivory, and porce-
lain copies were introduced by traders in the early twentieth century. Most, but
not all, dibadana have a hole through the centre. They were popular in the
1930s and many photographs from that period show women wearing them. In
the 1970s they were not seen in use but a few old women still had them in their
possession.
Ee
Fig. 115. Ear ornaments, dibadana, K68D, Klapwijk Collection.
(yo
LOBEDU MATERIAL CULTURE 18
x
Fig. 116. Woman and child. Note hair-style, ear ornaments and clothing, also
reed door of hut. Photo: E. J. Krige, 1936-8.
CLOTH
Machine-woven cotton cloth from India and Europe was brought into the
Lowveld by traders following the well-established routes from the port of Dela-
goa Bay to the interior. Striped salempore gained early acceptance among the
Lobedu as it did among the Venda who termed it nwalukambu. German print
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
introduced by the mission became popular and has remained so among older
women. Plain black morina was also an early trade cloth and it is still worn by
conservative Lobedu women. In the 1970s morina and salempore wraps, dugu
yanogeni and dugu yahopaketa, worn around the waist and across the shoulders
(Figs 116-117) were often braided with coloured cloth at the edges and worn
with a cloak, seyi, of coloured cotton. Local tailors took orders for machine
braiding (Fig. 118). There was a wide range of cotton cloth available in the local
stores but salempore and morina were still in demand by women who had not
adopted modern dress.
Fig. 117. Young girl with partly shaved head wearing salempore cloth,
and child wearing bead necklet, grass armbands and metal anklets.
PhotosE ve Ktiees 193658:
LOBEDU MATERIAL CULTURE 185
Ee ie al
Fig. 118. Local tailor, Lebiya’s village, 1975.
Members of the malé6b6 cult wore distinctive wraps of red and white cloth.
They said that these colours were favoured by the spirits.
Cotton head-cloths, mothini, common in the 1970s, were not worn in the
1930s except among the Christian community. Most women shaved their heads
completely but some allowed their hair to grow and styled it in particular
fashions. Some young women favoured leaving a crown of hair on the top of the
head, others shaved the hair to form certain styles (Figs 116-117), or parted it in
a number of bands (Tyrrell 1968: 63, 65).
SALT
A small amount of salt was extracted locally from the leaves of a certain
plant that were dried, mixed with water, filtered and the filtrate boiled away
until the salt remained. Most of the salt used by Modjadji’s subjects came from
the salt-pans in the Soutpansberg to the north and from Tsonga territory to the
east and was acquired through trade. Salt from pans was extracted in a filtration
process that can still be observed at Sautini, a hot spring on the confluence of
the Klein Letaba and Mamostwapi rivers.
The process is similar to that recorded by Junod in the early twentieth cen-
tury. The salt-laden earth is filtered through large grass-lined filters (Fig. 119A)
and the filtrate is evaporated, leaving the salt crystals, which are shaped into
small mounds for trading (Fig. 119B). This salt is much preferred to the factory-
processed product. Salt is added to the many relishes that accompany the rather
bland unsalted porridge.
186 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 119. A. Salt-filter made of mopane bark and branches, Sautini, 1973. B. Small mounds
of salt drying over warm coals, Sautini, 1973.
LOBEDU MATERIAL CULTURE 187
SUMMARY AND COMMENT
The above section is concerned with materials that were not indigenous to
the environment and therefore had to be obtained through trade or by travelling
outside the area. The most important material in this category is iron.
Iron was essential to the subsistence economy in that the exploitation of
both natural and domestic resources depended on the use of iron tools. Iron was
needed for the production of the basic agricultural implement, the hoe, as well
as for weapons and tools on which the technology was dependent. The fact that
this essential resource was not found in the Lobedu area gave it a high social and
economic value. Long distances were travelled to obtain ore, and iron objects
were valued items of trade. Throughout the Lowveld hoes were used as media
of exchange in trade and in marriage contracts.
Before the local metal-working industry had been largely undermined by
the appropriation of ore sources and the large-scale introduction of factory-
made copies of indigenous products, iron was imported from the Phalaborwa
and Venda areas both in the form of ore to be smelted and forged locally, and in
the form of complete artefacts ready to be hafted for use. By the late 1930s the
smelting-process was no longer practised in the Lobedu area (Krige 1982 pers.
comm.) and by the 1970s it was not even possible to record the complex pro-
cesses formerly used. Forging iron objects, -thula tsipi, from any available pieces
of metal continued for much longer and is still practised on a small scale by a
few craftsmen (often Tsonga-speaking immigrants) using scrap metal and
modern tools (Krige 1982 pers. comm.).
In the 1930s objects made of locally smelted and forged iron were rare and
by the 1970s they were extremely rare. Iron tools and weapons that have not
rusted away completely have been kept for religious purposes as sacred thugula
objects believed to have links with the ancestral spirits. When beer is offered at
a family shrine, it is poured over the thugula object, which is either embedded in
the shrine or placed on it to represent its deceased owner and, by association,
the ancestors in general.
By the 1970s purchased metal tools such as knives, saws, axes, nails, files,
drills, hammers, and pliers, were widely used, especially in making non-
traditional furniture such as chairs and benches, and in construction-work.
Window-frames, doors and roof structures were made using modern carpentry
tools and techniques. Although from the late nineteenth century carpentry
skills had been taught to the Christian community at Medingen, these skills had
not been widely implemented among traditionalists. Huts did not have win-
dows, and the doors were either of reed or carved from a solid piece of wood.
As mud-brick construction replaced the older pole-and-daga method, the struc-
ture of huts was subject to further innovation in the form of modern door-
frames and windows. The construction of these fittings required the use of
modern tools.
Traditional tools, however, were appropriate to their specific functions and
bédwana adzes and leh6r6é gouges (usually with blades forged from scrap metal)
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
are still used for carving drums or utensils from a solid piece of wood. The
important and particular relationship of tool form to artefact is apparent in this
continuity of technique, despite the availability and concurrent use of modern
tools.
Copper, which came from ore sources in the Venda and Phalaborwa areas,
was not essential to subsistence activities but was valued as a material for
drawing wire and for making beads. As in the case of ancestral iron objects,
copper ornaments that pre-date their mercantile equivalents have become
sacred thugula objects. Also in this category are the very old glass beads, some
of which are believed to have been brought to the area by the founding ances-
tors.
Krige (1982: 32) observes that a marked characteristic of Lobedu
material culture is the lack of elaboration in dress and ornamentation. She
relates this to the fact that the decorative aspect of ornamentation is
_ secondary to the association of beads with the ancestor cult and to their
protective and healing powers. Secondly, she draws attention to the fact
that, in contrast to the practice among exogamous Nguni groups, dress and
ornamentation was not an important part of courtship as, following the pre-
ferred pattern of cross-cousin marriage, most people grew up knowing
exactly whom they would marry. Ornamentation was thus only partly dec-
orative in intent and consequently modern glass beads have had relatively
little popular appeal.
Similarly simplicity characterized traditional dress even after trade cloths
were adopted. The striped salempore, German prints, and black morina cot-
ton cloths, which came into use during the nineteeth century, seem to have
been accepted more for their practical advantages in a warm climate than
for their decorative qualities.
The replacement of skin clothing by cloth wraps and modern garments
meant that people were dependent on an imported purchased commodity for
their basic clothing requirements. The traditional cloth wraps required little
or no sewing but, as modern dress became popular firstly among the Chris-
tian community, there was a need for local tailors and dress-makers (Fig.
118). This involved the use of a whole range of other imported goods from
sewing-machines to needles and cotton.
In the 1930s, with the important exception of metal goods, the balance
between locally produced and imported material items favoured local pro-
duction. Material innovations were not widely accepted except by the Chris-
tian community, and the subsistence economy was still viable. By the 1970s
the transition from a subsistence to a money economy was clearly reflected
in many aspects of the material culture. In addition to purchased alterna-
tives for many traditional objects, motor cars, bicycles and a wide range of
brand-name products were ubiquitous. It is relevant to consider the resili-
ence of the traditional technology in the face of fundamental economic
change.
LOBEDU MATERIAL CULTURE 189
MATERIAL CULTURE AND SOCIAL CHANGE
The present study, undertaken in the mid-1970s, aimed not only to describe
the material culture but to analyse changes that had occurred during the 40 years
since the Krige Collection had been made. The post-war period coincided with
an unprecedented economic development of the area, growth of light industries,
improvements in roads and communications, and increased white settlement fol-
lowing the virtual eradication of malaria. The black population of the area in-
creased far beyond the carrying capacity of the land and the domestic economy
became increasingly monetized. It is against this background of changing social
and economic conditions that changes in the material culture will be viewed.
In discussion of these conditions the author is indebted to E. J. Krige (1975,
1981) for her analysis of social change among the Lobedu, as well as for exten-
sive personal interviews and correspondence over the duration of the study.
Without her analysis based on data from both the 1930s and the period 1960-80,
it would not have been possible to gain insight into the interrelated nature of
material and social change.
HISTORICAL, ENVIRONMENTAL AND SOCIAL CHANGE, c.1880—1980
The final decades of the nineteenth century were significant in Lobedu his-
tory. During this period conflict over land alienation and taxation imposed by
the Volksraad of the Zuid-Afrikaansche Republiek increased, and led to con-
frontation and eventual Lobedu submission to imposed territorial arrangements.
The fixing of the boundaries of the Modjadji Location in 1892 greatly reduced
the arable land and grazing available to Lobedu farmers. White farmers settled
along the Mokeetsi River and on the land south and west of the Location
boundaries toward present Tzaneen. This dispossession of land coincided with
the loss of cattle through diseases, the rinderpest of 1896 being the most serious.
Few cattle survived this and a subsequent drought further strained the subsis-
tence economy and gave impetus to labour migration which had occurred on a
small scale from as early as the 1860s (Delius 1980: 296). Taxes payable in cash
forced men to enter the labour market, which had far-reaching social and ma-
terial implications. European dress was soon adopted by most men and the ac-
quisition of expensive items such as guns, ploughs, bicycles or sewing machines
was an additional incentive to earning money.
At first men stayed away from their home villages for fairly short periods,
and the percentage of men absent from the reserve at any one time was rela-
tively low, but the length of time away and the number of migrants increased
steadily. In the late 1930s migrants formed approximately 37 per cent of the
adult male population, by the end of the 1970s the figure was higher than 65 per
cent (Krige 1981: 181) and, although figures are not available for the 1980s,
there is no doubt that the number of migrants far exceeds the 1970s estimate and
continues to increase and includes growing numbers of women. In the 1980s
population density is such that the vast majority of people have no land to culti-
vate and are entirely dependent on the earnings of migrants.
190 ANNALS OF THE SOUTH AFRICAN MUSEUM
From the turn of the century onward migrancy not only brought money into
the rural economy and created new material needs but changed the balance of
labour within the Lobedu domestic economy by reducing the number of male
workers. Unlike the division of labour in Nguni cattle-based economies, in the
predominantly agricultural Lobedu economy men participated actively in crop
production. The reduced agricultural work force was to some extent compen-
sated for by the use of the plough, rejected in its early heavy form but accepted
later in a lighter form (Krige & Krige 1943: 325). Hoe-cultivation, however,
continued in many places as much of the Modjadji Location is too steep for
ploughing; furthermore, the plough had certain disadvantages—it destroyed
contour walling, and was dependent on draught animals and on good rainfall to
soften the soil prior to its use.
As the number of migrants increased, women played increasingly active
roles in the productive economy by becoming involved in the structural aspects
of hut-building, the moulding of sun-dried bricks and the making of string, bas-
kets and mats formerly made mainly by men. The importance of women’s roles
in the economy as well as in social, political, and ritual organization minimized
the disruptive effects of migrancy and was a stabilizing factor in the material cul-
ture, both in terms of the technology and the demand for its products.
In the late 1930s there was virtually no labour migration of women and even
by the late 1970s, despite the increasing tendency for younger women to work as
daily and weekly labourers on neighbouring farms or in light industries, there
were still relatively few women migrants (Krige 1981: 155). In general the tech-
nical processes such as pottery and calabash-making, for which women were tra-
ditionally responsible, continued to be practised to supply domestic needs. In
the case of pottery, production exceeded subsistence requirements as many
skilled potters made pottery for sale as well as for their own use. This was one of
the ways in which women responded to the transition to a money economy. Far
from dying out, by the 1970s the practice of this traditional craft had increased
to meet new needs. It is noted, however, that although in the 1970s there was
widespread use of traditional technology in the domestic sphere, this was per-
petuated mainly by older women and by those living at the capital which is
notably conservative. Young educated women showed little interest in acquiring
or practising traditional skills and looked to employment outside the area.
Post-war economic change
During the decade after World War II the previously unhealthy Transvaal
Lowveld was opened up to intensive white settlement following the virtual eradi-
cation of malaria (achieved by the aerial spraying of insecticides). The conse-
quent development of the rich mineral and agricultural resources of the area and
the growth of industries ended a period of relative isolation for the local black
inhabitants. There was an unprecedented demand for labour in the newly estab-
lished factories, mines, and plantations. A network of new roads made possible
a new form of migrant labour, that of weekend commuting to and from the
LOBEDU MATERIAL CULTURE 191
reserves. Migrants returning at weekends with cash to spend brought much
needed money into the economy and also provided a welcome market for the
sale of home-brewed beer, which enabled village women to supplement the
wages of their husbands.
By the 1970s women had responded to the inevitable shift from subsistence
agriculture (resulting from shortage of land) by seeking ways of earning money
at home. Apart from brewing beer for sale, women were involved in other
money-making activities such as sewing, the moulding of mud bricks, the
plastering of huts and floors, and decorating of courtyard walls. By the 1980s
growing numbers of women had become wage-labourers on tea-plantations,
fruit-estates, and in light industry. These economic changes have had far-
reaching social implications (see Krige 1981: 153-155).
Mission influence
During the late nineteenth century Lobedu values and traditions were sub-
ject to the impact of mission doctrine. In 1881 Reuter, of the Berlin Missionary
Society, established a mission station at Medingen and started working among
Modjadji’s people. Mission teaching was opposed to polygyny, bridewealth pay-
ment, and cross-cousin marriage; it stressed free choice of marriage partners and
discouraged the extended family, thereby threatening the essential character of
Lobedu marriage and social structure (Krige 1981: 151). Although relatively few
people were converted, the mission had a strong educational influence and pro-
vided training in technical skills, especially in carpentry and building methods,
and encouraged individual enterprise and private ownership of property. By the
mid-1930s it was predominantly the small Christian community (less than 5 per
cent of the population) that had accepted western dress and was involved in
money-making activities to enable them to meet the costs of education and the
material requirements of their new way of life (Krige & Krige 1943: 320, 326).
Although early educational influences had some impact on values among a
small section of the community, belief in the power of the ancestral spirits was
not much undermined by the ideology of mission teaching. The institutionalized
rain and fertility cult with Modjadji at its centre continued to command wide-
spread observance of custom, reflecting the tenacity of ancestor beliefs and the
importance of rain in a subsistence economy primarily dependent on crops and
indigenous plant foods. Even converts to Christianity believed that Modjadji
had the power to make rain, albeit by the will of God (Krige & Krige 1954: 81).
Persistence of belief in the ancestors, in the power of medicines to control the
forces of evil, and in the fertility and rain cult was still evident in the 1970s but
the relevance of a fertility cult in an increasingly monetized economy was open
to question. 7
Pressure on environmental resources
Pressure on natural resources and on land in particular increased signifi-
cantly in the period 1930-80. This was primarily due to an unprecedented
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
increase in population following the implementation of National Party government
policies regarding influx control of blacks in urban industrial centres and on
white-owned farms. The population density more than doubled between the late
1930s and the 1980 census when the average density had reached over 200 per-
sons per square kilometre. Of necessity much of the land formerly cultivated
was taken over for settlement. Such was the population increase that many
households had no fields at all. By the end of the 1970s the whole population
was dependent on wage labour and on buying their staple food, maize-meal.
The productivity of the remaining farm-land has been reduced by soil ero-
sion caused by over-grazing and the cutting back of protective bush and forest.
Plant resources basic to the traditional technology have been seriously depleted
and long distances must be travelled to obtain the materials required for special
purposes such as initiation costumes.
The increasing demand for fire-wood and the extensive use of indigenous
~ woods, including marula, for the firing of mud bricks has had a harmful effect on
timber resources. In the 1930s marula trees were never cut for fire-wood as the
nutritive content of the fruit and kernels was so highly valued (Krige 1983 pers.
comm.). In times of grain shortage the marula season in February and March
brought welcome relief to the hungry and bridged the lean months before the
reaping of the grain harvest. The present trend reflects both the breakdown of a
subsistence economy and the depletion of the environment to the extent that the
basic need for fire-wood outweighs all other considerations.
Investigation of the material culture in the 1970s showed a number of
responses to diminished natural and domestic resources, but the reaction to
changing economic conditions was more complex than a simple substitution of
new objects for old or the introduction of commodities previously unknown.
The 1970s inventory of material goods included many of the objects recorded in
the 1930s by Krige, but this did not mean that the pattern of their use had not
changed. In some examples, such as mortars for stamping maize, the frequency
of use had decreased considerably, in other cases substitute materials such as
sisal had replaced the indigenous fibres. Certain traditional objects had become
obsolete in response to social change, while others were used only on occasions
of ritual significance. Some of the technical skills were still practised, as in the
case of pottery, while others had disappeared completely. It became clear that a
number of variable factors came into play in accounting for the observed
changes and continuities. These included social, economic, technical, and ideo-
logical factors, the interaction of which resulted in the particular form of the ma-
terial culture at the time of observation. From a closer examination of specific
examples of change seen in relation to these factors, it is possible to gain some
insight into the nature of change in material culture.
CHANGE IN MATERIAL CULTURE c.1940—80
During the 1930s when the Krige Collection was assembled, the Lobedu
domestic economy was still largely self-sufficient and there was widespread
LOBEDU MATERIAL CULTURE 193
knowledge of the natural environmental resources utilized in the technology.
During the 40 years under discussion, for reasons outlined above, there was an
inevitable transition to a money economy. Some of the ways in which this funda-
mental economic change affected the pattern of use of particular utensils and
structures is discussed below.
Granaries and utensils associated with harvest
In a grain-based economy full granaries may be seen as indicators of pro-
ductivity and symbols of prosperity. In the 1970s, however, the disuse of grana-
ries reflected the converse. According to Krige (1982 pers. comm.) the 1930s
situation was strikingly different. Every homestead had kheasho or ledulu (see
p. 142) granaries for storing maize on the cob and many also had an under-
ground pit, kheledese (see p. 140), in the cattle-kraal or in the khdér6é for the
long-term storage of the decobbed maize. Large khesého grain baskets (see
p. 74) for more accessible grain storage were widely used and at harvest-time
temporary storage structures, moako and letole (see p. 143), for millet and sor-
ghum were set up in the fields prior to threshing (see Fig. 53). By the 1970s crop
yields could not satisfy basic grain requirements and there was no surplus to be
stored. The few granaries observed by the author were empty and in disrepair;
grain-pits had become obsolete.
Also prominent in the 1930s was the large mothatha basket (see p. 69)
used for carrying the harvest from the fields. As harvests declined, so did the
need for this basket, added to which was the fact that enamel basins proved use-
ful substitutes when carrying-utensils were needed. None the less, in the 1970s
mothajtha baskets were still occasionally seen in use in cooking-huts and court-
yards for holding maize-meal or other foods and, in areas where cultivation con-
tinued, they were still made prior to the harvest to supply a seasonal demand.
In the 1930s harvests were large enough to require transport by wooden
sledges. Fairly widely used after their early adoption from white farmers in the
Lowveld, these triangular wooden sledges were no longer in use in the 1970s.
Considering the soil erosion that they caused their use had been discouraged by
agricultural extension officers but, in addition, the diminished harvests rendered
them unnecessary.
Utensils associated with the hand-milling of maize
Ubiquitous in the 1930s, the wooden mortar, Jefudu, and pestle, mose (see
Fig. 52), were used two to three times a week throughout the year, the pre-dawn
thud of stamping being one of the characteristic village sounds (Krige 1983 pers.
comm.). By the 1970s the use of mortars and pestles for producing maize-meal
was limited to a few months after the harvest. Thereafter most people were
dependent on buying their staple food and the mortar and pestle were seldom
used.
With the reduced domestic stamping of maize, the varied dietary pattern
associated with different textures of meal produced at the successive stages in
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
the process of stamping (see p. 78), was largely replaced by a monotonous and
less healthy diet based on highly refined, bought maize-meal.
Also related to the decrease in frequency of stamping maize was the decline
in the use of the khegdy6 mat (see p. 84) used for drying the fine meal, and the
wide-mouthed /ebéda pot (see p. 66) used for steeping maize during the stamp-
ing-process.
Interestingly, the /esél6 winnowing-basket (see p. 74), widespread in the
1930s for use during the stamping-process, remained in regular use not only for
winnowing when necessary but for a number of other domestic uses. It served as
a general-purpose kitchen utensil, a tray, a lid to cover the mouth of a pot, a
base for drying relish, or for holding and carrying garden produce. This multi-
purpose utility seems to have ensured a continued demand for the lesél6.
Utensils associated with cooking and the mosha
Although there was a widespread transition from home-grown to bought
maize, maize-meal still remained the basis of the diet and the manner in which it
was prepared and served remained relatively unchanged. Most of the traditional
domestic utensils associated with cooking were still much in evidence even in the
early 1980s. The khelalél6 cooking-pots of various sizes and morifhi bowls (see
p. 64) were used in almost every household for preparation of the daily meal, as
were the wooden spoons, maf6é (see p. 110), and stirrers, mafhéthd (see
p. 111), used at different stages during the cooking of vhoswa, the staple maize-
meal porridge. Most cooking still took place on an open hearth in the mosha or
inside the cooking-hut, the cooking-pot being supported by three suitably pos-
itioned stones or, in some cases, by a purchased triangular metal stand known as
a ‘drievoet’ (‘three-foot’). Having been prepared in the traditional manner, the
porridge was served into well-used and often carefully repaired calabash vessels,
zwithéba (see p. 153). The continued use of these utensils reflects the value
placed on them by village women. Although used side by side with enamelware
and plastic utensils, many of the locally made domestic utensils were preferred
for their particular qualities and usefulness. Thus the pot, modzéha (see p. 60),
used for storing drinking-water at home had not been replaced by tin or plastic
containers because the porous, cooling quality of the earthenware made it pref-
erable to alternative containers. For transporting water, however, lighter and
less breakable containers have advantages over pottery and consequently, since
the 1930s, tins and plastic buckets had entirely replaced pottery vessels for this
purpose.
In addition to particular material properties that give certain utensils practi-
cal advantages, characteristic posture while working is a pertinent factor when
considering the usefulness of particular objects or substitutes. Thus the hand-
broom lefielo (see p. 89), has not been replaced by the long-handled European
broom because it is more convenient to have at hand for sweeping while cooking
at the open hearth. Furthermore, as Krige has observed (1983 pers. comm.),
Lobedu women do not see bending down as a disadvantage because most
LOBEDU MATERIAL CULTURE 195
activities of the mosha are carried out at ground level in kneeling or bending
position.
Earning money at home: beer-brewing for sale
The transition from a subsistence to a money economy deprived village
women of their economic self-sufficiency. In most cases their husbands’ wages
were insufficient to meet the basic needs of food and clothing. A response to this
situation was that women became increasingly involved in money-making activi-
ties at home. Whereas in the 1930s it was mainly the Christian community that
made ‘a business’ of sewing or selling pottery, by the 1970s an informal sector of
the cash economy was developing rapidly. One of the most successful ways in
which women could earn money was to brew beer for sale.
A number of changes in material culture were related to the change from
brewing beer largely for social or hospitality purposes, as was the case in the
1930s, to the more recent brewing of beer for sale, mainly to supply weekend
commuters and visitors (access being facilitated by improved roads and trans-
port). The increase in scale of brewing demanded larger utensils—the large clay
nkhoé (see p. 60), traditionally used in brewing beer, was replaced by the even
larger “44 gallon’ drum, and the woven beer-strainer, leth6d6 (see p. 86), was
replaced by a capacious wooden-framed wire-gauze sieve, sef6 (see p. 114). As
neither the traditional beer-strainer, nor any commercially sold strainer was big
enough to cope with very large volumes of beer, enterprising craftsmen devel-
oped the sef6 (term taken from Afrikaans ‘sif’) to meet the needs of large-scale
brewing. It is an interesting example of a borrowed idea modified to suit specific
requirements.
By the 1970s, in addition to the time-honoured pattern of sharing beer
among kinsmen and neighbours on occasions of social or ritual importance, an-
other form of social beer-drinking had developed in response to the selling of
beer. In the older ‘hospitality’ beer parties the utensils used were the traditional
beer-pots (see p. 62) and calabashes. The long-handled calabash khehé kha
byalwa (see p. 153) in which beer was served was passed freely from person to
person, showing generosity and bonding social ties. The etiquette of the oc-
casion was dignified and ordered, the formality being expressed in the conven-
tional seating of men and women on either side of the server and in the use of
traditional utensils. By contrast, social occasions which centred around the sale
of beer were informal and traditional utensils were inappropriate. The need for
containers of standard volume in which the beer could be sold to individuals was
met by glass jars or tins of regular size. Instead of the beer circulating among a
group of friends, the buyer tended to consume the beer for which he had paid
and, as a customer, his behaviour was not subject to the etiquette appropriate to
a guest. Thus the difference between these two forms of beer-drinking, both of
which occurred in the 1970s (the conventional form being rare and usually as-
sociated with ritual occasions), was clearly reflected in the related utensils and in
the pattern of their use.
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
The effect of migrant labour on the use of certain utensils
As noted previously, the migrant-labour system brought money and a wide
range of imported commercial goods into the domestic economy. It also had di-
rect and indirect implications for the manufacture and use of certain traditional
utensils.
A direct response to the decrease in the number of men in rural villages was
an increased participation of women in craft activities formerly undertaken
mainly by men. This trend, which was noted in the 1930s in relation to hut-
building and the manufacture of mats, certain baskets and cordage, continued in
the following decades. Where demand for a product continued, as in the case of
sleeping-mats, women became the main producers both for domestic use and for
informal sale.
One of the crafts, however, that remained exclusive to men was wood-
working. Women were unaccustomed to using carving-tools and the arduous
_ work of felling trees or cutting large branches, often deep in the bush, was more
appropriate to men. Furthermore, the rough carving of large objects often took
place in the bush to avoid having to transport the heavy timber. Although
migrancy reduced the potential number of craftsmen, there always had been
some degree of specialization and enough carvers remained to supply domestic
needs. Many of the carvers active in the 1970s were older men or those unable
to find employment in town, who could earn some money locally by making
wooden utensils for sale. Carpentry skills, which had been taught at the mission
since the late nineteenth century, were in demand for the construction of new
houses and other buildings within the reserve. Thus both traditional wood-
working technology and introduced carpentry techniques were utilized to supply
specific demands.
The direct effect of migrant labour on the customary pattern of marriage
(see Krige 1981: 148-157) had indirect effects on the material culture by chang-
ing the social context in which certain objects had been of practical and symbolic
significance. The decline in use of the kherodwana serving-basket (see p. 80) is a
good example of a utensil that became obsolete as an indirect result of migrant
labour. It was customary practice for a bride to take a kherodwana to her new
home as a symbol of willingness to serve and honour her husband. In this
basket, covered with a lid, she placed the daily meal that she prepared for her
husband. As it became increasingly common for marriages to be contracted
while the groom was working in town, the need for this special-purpose basket
declined. By the 1970s it was no longer in use at all and only a few older men
could recall the intricate way in which it had been made.
Material expressions of beliefs and values
In the context of ritual, objects and the materials from which they were
made were invested with symbolic meaning and their use was prescribed by cus-
tom. For example, the vuhwera and vyali costumes (see p. 93), made from
materials associated with rivers or damp places, were believed to enhance fertility
:
LOBEDU MATERIAL CULTURE 197
and prosperity. The choice of materials was based as much on a complex system
of values as on technical considerations. Similarly, river stones and certain
plants were believed to have ‘cooling’ properties and to embody positive, pro-
tective forces. The material dimension of ritual in the form of medicines, thugula
objects, shrines, certain living plants, or offerings of beer made from Eleusine
formed part of a much wider cognitive system that communicated meaning to
those who understood the symbolic associations.
In order that meaning could be understood clearly, material symbols had to
be conventional and not subject to idiosyncratic variation. The effectiveness of a
non-verbal language depends on the continuity of the symbolic associations. In
the Lobedu case there was evidence that knowledge of the symbols necessary to
communication through ritual action and the associated objects remained alive
even in the 1970s. This could be accounted for by the continued observance of
customs, by the tenacity of ritual practices and by the importance of women in
the sphere of ritual. In the long term, however, school education and the conse-
quent change in values cannot but undermine the relevance of a traditional
world view.
CONCLUSIONS REGARDING THE NATURE OF CHANGE
From the examples outlined above it is clear that an interplay of economic,
social and ideological factors accounted for change and continuity in the ma-
terial culture and that the nature of material change was more complex than
simple replacement of hand-crafted artefacts by machine-made substitutes. Dif-
ferent kinds of change related to particular circumstances and social contexts—
by the 1970s some artefacts, such as granaries, had disappeared completely in
response to economic change; others, such as calabash drinking-vessels for beer,
were retained for traditional social or ritual purposes, whereas modern substi-
tutes were used when beer was sold; the physical properties of certain utensils,
such as earthenware pots for storing water, made them preferable to substitutes,
thereby ensuring their continued use, and the multi-purpose utility of some
objects, as in the case of winnowing-baskets, made them less likely to become
obsolete than utensils of more restricted utility.
Early changes in dress related to mission and school influence as well as to
the requirements of town life. Social and religious distinctions were expressed
outwardly in differences in style of dress and, in some cases, in colour and type
of cloth. The expression of consciousness through clothing or ornamentation
continues to adapt to changing circumstances. In the 1980s a trend toward ‘neo-
traditional’ dress, arising from national consciousness, was apparent among edu-
cated women. This was particularly evident at the coronation of Modjadji V in
1982 (Krige 1983 pers. comm.).
Numerous examples confirmed that change in material culture could only
be understood with reference to the wider social and economic context. The
undermining of the metal-working industry that occurred before the period
under discussion was a direct result of colonial appropriation of the ore sources
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
and smelting-sites, concurrent with the large-scale importation of cheap iron-
ware from Europe. In addition to the fact that the termination of local produc-
tion forced people to buy alternatives was the fact that many of the imported
tools, such as knives, axes or saws, had practical advantages both in terms of
availability and function. The saw, saha, for example, had no traditional equiva-
lent and met a real need; imported knives were very efficient and easier to
handle than the older mofhago. There was not, however, a total replacement of
old tools by new. Traditional adzes and gouges (see p. 101) were still the most
appropriate tools for carving certain wooden objects and the demand for them
remained. From the 1930s onward this demand was met by the forging of iron
(-thula tsipi) from scrap metal, often undertaken by Tsonga-speaking immigrants
from the Phalaborwa area (Krige 1983 pers. comm.). Thus, although in the
1970s there was an apparent continuity in the use of certain tools, the method of
their manufacture had changed considerably.
In the case of pottery the technology had remained remarkably constant
but, in response to changing economic conditions, the scale of manufacture had
increased far beyond subsistence needs. Conversely, the decline of skin-working
presents an example of diminished production linked to changes in clothing cus-
toms, decreased demand for articles made of skin, and consequent decline in the
related technology.
Basketry, in many respects, seemed to reflect a continuity of use and manu-
facture, but on investigation it turned out that women had become increasingly
involved in the making of certain baskets and mats in response to the reduction
of craftsmen caused by migrancy. An unqualified observation of continued use
of these items might have masked the significant social change that underlay the
increased participation of women in this form of production.
In view of the fundamental economic changes that have occurred over the
past 40 years it is perhaps surprising that so many of the traditional technical
skills have remained useful and relevant. Women in particular have proved both
conservative and adaptable in this changing situation by using traditional skills
to generate income and meet new economic needs. It is predictable that, as
increasing numbers of women enter the labour market, the previously conserva-
tive influence of women will be reduced.
In attempting to gain some understanding of the nature of change in
Lobedu material culture the limitations of a purely technological approach became
apparent and it was fortunate that a wealth of anthropological data, collected
and analysed by E. J. Krige, could be drawn on to inform the discussion of
change. The general conclusion that can be drawn from this study supports the
premise that change in material culture is inextricably bound up with social and
economic factors and with ideological values; the precise nature of the links and
the way they articulate depend on particular circumstances and are part of a
continuous process of change.
LOBEDU MATERIAL CULTURE 199
ACKNOWLEDGEMENTS
Part of the data published here formed part of a Master’s thesis in Anthro-
pology under the supervision of Professor G. M. K. Schitler, University of Stel-
lenbosch.
My warmest appreciation goes to Professor E. J. Krige for her assistance
throughout the study and for her constructive comments on the manuscript; to
Mr Simeon Modjadji and the people of Modyjadji’s village for their willing
co-operation; to Miss E. M. Shaw for her comments on the manuscript and
permission to consult her unpublished manuscript on basketry; to Dr N. J. van
Warmelo for advice on the vernacular terms; to the Compton Herbarium for
identification of botanical specimens; to Mrs June Hosford for photographic
work; to Mr V. Branco for the line drawings, and to Mrs Marcelle Scheiner for
typing the manuscript.
The Department of Co-operation and Development and the Lebowa Govy-
ernment are thanked for permission to work in the area. Financial assistance
from the Human Sciences Research Council is gratefully acknowledged. Opin-
ions expressed and conclusions drawn are those of the author and should not be
regarded as representative of the Human Sciences Research Council.
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6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
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Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15A
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Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
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PATRICIA DAVISON
LOBEDU MATERIAL CULTURE
A COMPARATIVE STUDY
OF THE 1930s AND THE 1970s
PART 4 SEPTEMBER 1984 ISSN 0303-2515
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(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 94 _ Band
September 1984 September
Part = 74 Deel
MORPHOLOGICAL AND BIOLOGICAL NOTES
ON SOME SOUTH AFRICAN ARTHROPODS
ASSOCIATED WITH
DECAYING ORGANIC MATTER
PART 3
THE FAMILIES DERMESTIDAE,
CANTHARIDAE, MELYRIDAE,
TENEBRIONIDAE, AND SCARABAEIDAE
(COLEOPTERA)
By
A. J. PRINS
Cape Town Kaapstad
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MORPHOLOGICAL AND BIOLOGICAL NOTES ON
SOME SOUTH AFRICAN ARTHROPODS
ASSOCIATED WITH DECAYING ORGANIC MATTER
PART 3
THE FAMILIES DERMESTIDAE, CANTHARIDAE, MELYRIDAE,
TENEBRIONIDAE, AND SCARABAEIDAE (COLEOPTERA)
By
A. J. PRINS
South African Museum, Cape Town
(With 24 figures)
[MS accepted 2 April 1984]
ABSTRACT
The immature stages of twenty species of polyphagous beetles belonging to the series
Bostrychiformia, Elateriformia, Cucujiformia and Scarabaeiformia were collected along a nar-
row strip between Mossel Bay and Elands Bay and in small areas around Laingsburg and
Tulbagh. Their morphology is illustrated and their biology and association with decaying organic
matter are noted. Observations on some families of minor importance are also included. Most of
the species discussed are directly involved in the breakdown of organic matter.
CONTENTS
PAGE
Introduction and observations on minor families............ 204
Families Coccinellidae and Chrysomelidae............. 205
Families Elateridae and Anthicidae................... 206
Families Ptiliidae and Endomychidae ................. 206
amilyiClenidacr Wii ers Sook te ince ee ae ana ae 206
Families Colydiidae, Cucujidae and Ptinidae ........... 208
RamilyaNitidulidacie «25 Avi i. See 8 a Seinen eee 209
Families Cerambycidae and Curculionidae............. 209
Morphological descriptions and biological notes ............ DAZ
amilyMenmestidaer 2.4 1e steerer. oe eee tes INDE
Dermestesimaculats de Geena sens ne een DAS
DermestesiaremdeGect tas heme a orc ZA)
Dermestes peruvianus Castelnau.................. DOM
anmlyaCanth anda’ Atatt re eer re es ae 228
Avjronyeha picta (Wicdeniann))sa. ane sees eee 224
army NMC lw ae age es ce vee seu ayer tatnsy: taps acy 226
IMelyrisnvinidis i abniCiUS)}.4 324. eee oe cere D2
Familygkenebmomidacre, sy scs co. soon spoon ee JEM |
Gonocephalum arenarium (Fabricius) ............. 258
Pachyphaleria capensis (Castelnau) ............... 236
Tribolium destructor Uyttenboogaart ............. 239
Alphitobusidiaperinus (Ranzen)..5 2.445220" 20 en 243
203
Ann. S. Afr. Mus. 94 (4), 1984: 203-304, 24 figs.
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
PAGE
Fanuly Scarabacidae ta seca. ce cee ee es 249
Apnodmsilacius Wiedemann ae eee juss)
Apnodiusimoesius Fabricius #225944. 6e sale ee 263
VAD NOGIUSIPLOCERUSIAatO dhe ae ran ere 265
Aphodiusiconsimilis Boneman — 4-4 4e.45 ee nee 269
Onis aygidus (Fabricius): 22... -e.4606 sae nee ae 272
Oniucellus picms(Hausmann))).9)..522 2.5505. 555- 274
Oniticellus planatus Castelnau.................... 278
Rachnodasinuata(Habticius)ia se eae aan 281
Rhinocoeta cornuta\(Fabricius)).....)..5...4. 25900 285
Temnorrhynchus retusus (Fabricius) .............. 290
roxy paroideslaroldin eee ne 296
ACKn owled eemiemtse es nea ricm rt Nn celts one eeu te rca 300
IRCICRENCESIS hs aati d casas hes surance ehh, ale, Sadar age a 300
AND DEC VIAUONS¥e 7 one, Sei erate Rete reer eee cat on ane 302
INTRODUCTION
AND OBSERVATIONS ON MINOR FAMILIES
During the surveys of arthropods associated with decaying organic matter
thirty-four species of beetles were collected in a narrow strip along the southern
and western coastal regions of South Africa and also a small strip between
Laingsburg and Beaufort West as well as between Worcester and Tulbagh.
Fourteen species which are mostly predacious were discussed in the second part
of this series; the immature stages of the remaining twenty polyphagous species
belonging to the series Bostrychiformia, Elateriformia, Cucujiformia, and Scara-
baeiformia are treated here.
The Scarabaeiformia include the coprophagous, phytophagous, myrme-
cophilous and termitophilous scarabs, the termitophilous valgines as well as the
myrmecophilous clambids, and two other small families, the helodids and dascil-
lids. The Elateriformia, Cucujiformia, and Bostrychiformia contain species of
diverse habits and all have members that are of economic importance, being
either entomophagous, phytophagous, xylophagous, or pests of stored products.
The curled-up scarabaeiform larvae of the cockchafers, dung-rollers, fruit-
beetles and monkey-beetles are common in most soils containing plant roots or
decaying organic matter. Predacious species, however, such as ladybirds and
leather-winged or soldier-beetles (family Cantharidae) have caraboid or cam-
podeiform larvae, whereas the eruciform types of the skin-and-hide beetles and
those of melyrids and clerids are often found in dried vegetable or animal matter.
The trogositids (family Trogositidae) (Fig. 13C), which are near the clerids as far
as larval characters are concerned, are often attracted to carcasses and cadavers
during the later post-mortem stages of decay, particularly after rains.
The adult beetles are very often gregarious or congregate in large numbers,
such as the orange and black-coloured ladybird, Lioadalia flavomaculata
(de Geer), of which vast numbers are occasionally found on fresh and semi-fresh
kelp strings in the supratidal and also in the intertidal zones during the early
summer. There is no explanation for this behaviour, but it perhaps coincides with
SOUTH AFRICAN ARTHROPODS 205
a shortage of its main food supply, aphids, at this time of the year, and they are
then forced to search for new resources. Masses of dead and dying ladybirds,
mainly the predacious L. flavomaculata, together with Adonia variegata (Goeze),
Cheilomenes lunata (Fabricius), and Scymnus moreletti Mulsant, and various
other beetles and stink-bugs are often found on the intertidal zone along the west
coast. These insects have been blown out to sea by strong winds and are then
washed on to the beaches.
Various beetles, some of which may be of economic importance, are
attracted to dry or decaying organic matter; they may play an important role in
the destruction of such material, or they may merely utilize it as shelter (e.g.
under large dry cow-pats), particularly in open veld with little vegetation.
In the cases mentioned below, no morphological data on the larvae could be
compiled. The adults of most of the species, however, were in some way associ-
ated with, or are related to, species attracted to organic matter.
Families Coccinellidae and Chrysomelidae
Apart from the above-mentioned ladybirds, another very small predacious
species, Cranophorus varius Weise, which, according to collection data, is
endemic to the Cape Province, was attracted to carcasses where a second and
third sarcophagous stage occurred after repeated rains during the late autumn and
early winter. It is about 1,9 mm long, shiny brown to piceous and covered with
short fine hairs.
The adults of the phytophagous ladybird Epilachna andulata Thunberg, and
of the red and black-striped leaf-eating beetle Chrysomela fasciata de Geer, are
sometimes found to shelter under semi-dry cow-pats. Adults and larvae of both
species feed on the leaves of plants of the daisy family (Compositae) (Skaife
1953). C. fasciata often causes extensive damage to foliage of Arctotheca populi-
folia on the sand-dune system along the south-western coastal belt during late
autumn and early winter. All the developmental stages of E. andulata may at
times be collected on the same plant.
Pupae of another phytophagous ladybird, Henosepilachna elaterii (Rossi),
were occasionally collected under semi-dry cow-pats along the coastal areas
during autumn, together with adults of E. andulata. Henosepilachna elaterii were
often associated with the black and orange-coloured H. gibba (Thunberg) in the
Saldanha— Vredenburg area, also under cow-pats. The pupae of H. elaterii are
yellowish with blackish spots, and the adult beetles, which emerged during March
and April, are 5,8—8 mm long, cadmium yellow, with large oval black spots.
The larvae of some beetles such as certain halticids or flea-beetles (family
Chrysomelidae) contain a toxalbumin, which may cause severe pain and even
death in warm-blooded animals. Death by paralysis is apparently caused by a
lethal saponin in the blood of the larvae of the bushman-poison beetles, Diamphi-
dia and Polyclada spp. (Shaw et al. 1963). These are all yellowish beetles with
larger or smaller blackish spots or marks on the elytra, and they vary from 10 to
15 mm in length. The larvae of the carabids of the genus Lebistina, particularly
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
L. peringueyi Liebke, L. holubi Peringuey, and L. subcruciata Fairmaire, which
are ectoparasitic on the larvae of the above-mentioned poison-beetles, are even
more poisonous than their hosts (Koch 1958). These carabids, about 15 mm in
length, are also yellowish beetles, some with oblong blackish markings on their
elytra. A few of these parasites and their hosts have been observed by the author
under semi-dry animal droppings.
Families Elateridae and Anthicidae
Orthosomatic larvae are found in many families of which the adult beetles
are associated with decaying organic matter; only in a few instances, however,
have the larvae been observed in this medium. Certain click-beetles (Elateridae)
such as Lacon amplicollis Boheman, a brownish to blackish species about 6,6 mm
long, and some Cardiophorus spp. are often found in the veld under dry cow-pats
which do not harbour carabids. Larvae of these beetles, also known as wire-
‘worms, are elongate and cylindrical with a prognathous head, and may be
predacious, phytophagous, and even liminous, as is the case with certain can-
tharids. Ant-beetles (Anthicidae) are common under dry cow-pats almost
throughout the year, particularly the dull black Anthicus stygius la Ferté, and the
larger shiny black Formicomus caeruleus (Thunberg) (about 4,2 mm long). Only
one species, the shiny, dark-brown, wingless Anthicus apterus van Hille
(Fig. 8M), was observed on carcasses during the ceratophagous stage of decay,
but it was never seen on animal droppings in the veld.
Families Ptiliidae and Endomychidae
One of the most common beetles found in semi-fresh to almost dry dung is a
black unidentified species of ptiliid, about 0,8 mm long. These insects, easily
recognized by their feather-like wings, are some of the smallest of beetles known.
Their campodeiform larvae resemble those of the rove-beetles in shape as well as
in the pseudopodium-like tenth abdominal segment. They are active and are
often found together with the adults even on compost heaps and decaying kelp on
the beaches. According to Dybas (1976) the larval mouth-parts are not adapted
for a carnivorous mode of life and they apparently feed on the organic matter and
on fungus spores as in the case of Mycetaea ovulum Wollaston (Endomychidae).
The latter is a small, oval, shiny brown fungus-beetle (about 1,3 mm long) often
found in association with this feather-wing beetle, and which is, according to
collection data, endemic to the Cape.
Family Cleridae
The chequered or clerid beetles are elongate, sombre coloured to metallic
greenish or bluish or even brilliant red and yellow. Elytra are usually entire and
tarsal formula is 5—5—5. The larvae are orthosomatic with a prognathous head;
epicranial halves are separated by a gular area as in melyrids and the molar area
of mandibles is lacking; dorsolateral glands are apparently absent in members of
this family.
SOUTH AFRICAN ARTHROPODS 207
About 300 species have been recorded from the southern African subregion.
Most of them are predacious in both the adult and larval form and are therefore
beneficial; only three species, as far as is known, have become pests of stored
products with a high fat content. Of these, two cosmopolitan species, the red-
legged ham-beetle, Necrobia rufipes (de Geer), and the red-shouldered ham-
beetle, N. ruficollis (Fabricius), occur in South Africa. Both these beetles are
attracted to decaying carcasses of land and sea mammals and human corpses
during the casein fermentation period. They are both long-lived and survived in
the laboratory from March to January or February (a period of 10—12 months),
copulation being observed during October. Simmons & Ellington (1925) give the
longevity as 14 months. Larvae of both species were found to prey on the larvae
of the skin-and-hide beetle Dermestes maculatus de Geer, and those of the
cheese-skipper Piophila megastigmata McAlpine.
Necrobia rufipes is a metallic greenish-blue beetle (4,4—6,6 mm long) with
light reddish legs. The eggs are banana-shaped, yellowish white, and those
collected on a decaying turtle measured 1,10 by 0,24 mm. The incubation period
in the laboratory (25-26 °C) was 2-3 days and the life-span of five larvae occu-
pied 90-272 days on dried beef, the pupal period being 9-13 days (21-25 °C).
According to Simmons & Ellington (1925) the shortest developmental period is
30 days, which includes 17 days as growing larva and 13 days within the cocoon,
until emergence of adult (daily mean temperature 24,4 °C).
Necrobia ruficollis, on the other hand, is somewhat smaller (4,5-5,5 mm
long, blackish blue in colour with reddish pronotum and elytral shoulders, and
also with reddish legs. Eggs are similar to those of N. rufipes; those collected on a
seal carcass measured 0,88 by 0,28 mm. The incubation period in the laboratory
(21-25 °C) was 2-5 days, the larval life-span being 104—203 days under similar
conditions as those of N. rufipes. In both species a papery cocoon is produced by
the larva in which pupation occurs. The pupal period occupied 9-14 days.
Descriptions of the larva and pupa are given by Heeger (1848) and Scott (1919),
and a general description of a clerid larva by Peterson (1967).
Both the above species as well as the smaller Opetiopalpus collaris Schonherr
(3,6-4,4 mm) and Corynetinus fimetarius Wollaston (3,2-3,9 mm) have been
collected under semi-dry to semi-fresh cow-pats on the beach and further inland.
Both these latter species are blackish blue, the first with reddish and the second
with bronzy pronotum.
In Cape Town at least four species of clerids have been recorded in timber,
mostly Eucalyptus and kiaat, with the adult beetles feeding on the larvae of
powder-post beetles (Lyctus species). The largest of these is Cylidrus fasciatus
Castelnau (7,9—9,2 mm long), a somewhat shiny dark-brown beetle with a fairly
broad yellowish transverse band across the middle of its almost impunctate elytra.
It is widely spread in the Subsaharan and Malagasy regions (Corporaal 1950).
Paratillus carus (Newman), the white-banded clerid, which is of Australian
origin, has piceous elytra with a very narrow whitish-yellow transverse band, the
head and pronotum being reddish brown. It is slightly smaller than C. fasciatus
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
(5,3-6,1 mm long). Its adults are often observed in the Cape Town docks in
Japanese oak, where it feeds on both adults and larvae of powder-post beetles. It
has also been introduced into England (Winkler 1960). According to Fisher
(1944) it is a common predator on Lyctus beetles and is also one of the most
numerous and active predators upon larvae of the smaller timber-borers in
Australia (Froggatt 1927).
Pallenis misella Boheman (4,9 mm long) is dark brown with a narrow yellow-
ish transverse band over the punctate elytra, but it differs from the other three
species in the pronotum, which is deeply and widely excised on each side of its
posterior part. It is widely distributed in South Africa, including records from
Natal.
The cosmopolitan small white-blotched clerid, Tarsostenus univittatus
(Rossi), is of similar coloration to P. misella, but has a narrower yellowish
transverse band on the punctate elytra and is smaller (3,2—4,6 mm long); it is
_ widely distributed in South Africa as indicated by collection records from the
Sandveld along the west coast and from Transvaal. According to Laing (1928) its
adults appear at almost the same time as the adults of its host. Froggatt (1927)
obtained this species from wood infested with larvae of Heterobostrychus aequalis
Waterhouse.
A metallic-blue clerid Cylidrus wallacei Thomson, almost 12 mm long, was
found in Bellville, Cape Province, during March 1983 in Koompassia malaccensis
logs imported from Malaysia, and which were infested with larvae and adults of
the yellow-legged auger-beetle, Xylothrips flavipes (Illiger) and the previously
mentioned shot-hole borer, Heterobostrychus aequalis. Both bostrychids are
widely distributed in the Far East (Froggatt 1927).
Trichodes aulicus Klug, another beautiful metallic-blue species about 8 mm
long, but with three orange patches on the elytra, was bred during October from
the nest of a small megachilid bee in the Calvinia district. According to collection
data it is widely spread in the north-western Cape Province; otherwise its habits
are unknown.
Families Colydiidae, Cucujidae and Ptinidae
The cylindrical bark-beetles (Colydiidae) and the flat-beetles (Cucujidae),
particularly Euxestus phalacroides Wollaston and Monotoma spinicollis Aubé,
which are common during the last or ceratophagous stage of decay of carcasses,
were observed to be very numerous in dry cow-dung along the west coast of the
Cape Province. They were found to feed on the dry particles and were often
associated with the shiny black indigenous spider-beetle, Pseudomezium coc-
quereli (Fairmaire), which is about 2 mm long. Both the first-mentioned colydiid
and cucujid are widely distributed in South Africa and overseas (Hetschko 1930a,
1930b) and are small (1,9—2,3 mm long). Euxestus phalacroides is oval and shiny
brown. Monotoma spinicollis is elongate and dull dark brown with serrate lateral
prothoracic margins.
Spider-beetles (Ptinidae) are common in the dry decay stage of carcasses and
SOUTH AFRICAN ARTHROPODS 209
cadavers, and at least four cosmopolitan species occur in South Africa, of which
Mezium americanum Castelnau and Gibbium psylloides (Czempinski) are almost
glabrous and shiny. The latter, also known as the storehouse-beetle, usually
occurs in ships carrying dried foodstuffs and is often reported as damaging maize
kernels under such conditions. This species, together with the drugstore-beetle,
Stegobium paniceum Linnaeus, and the tobacco-beetle, Lasioderma serricorne
Fabricius, has been found in the tomb of Tutankhamun (Hinton 1945). The other
two cosmopolitan ptinids, Tipnus unicolor (Piller & Mitterpacher), found in
walnuts, and Pseudoptinus lichenum (Marsham), introduced from Europe, are
hairy. Larvae of P. lichenum are known to bore in dry wood and bark of fig trees
(Hinton 1941).
Another small, brown, hairy indigenous spider-beetle, Stethomezium
squamosum Hinton, was found in the roots of the shrub Maerua racemulosa,
which are eaten by the local people during food shortage; this shrub occurs from
the eastern Cape Province to Swaziland. Hinton (1943) also records Ptinus tectus
Boieldieu together with other beetles in the roots of this plant, as well as Mezium
natalense Peringuey breeding in blood-meal in Zimbabwe. A rather small,
unidentified, shiny-brown Meziomorphum sp. covered with fairly long spinose
hairs and which is about 1,9 mm long, has been observed to feed on owl pellets in
the vicinity of Cape Town.
Family Nitidulidae
Various undetermined sap-feeding beetles often appear on decaying animal
matter, mostly on carcasses during the late dermatophagous and early cerato-
phagous stages of decay. Two well-known members of this family, the dried-fruit
beetle, Carpophilus hemipterus (Linnaeus) and the corn-sap beetle, C. dimidiatus
(Fabricius), are, according to collection data, widespread in South Africa. Carpo-
philus hemipterus is dark brown with large brownish-yellow marks on the elytra
and was common in soil where fish-meal was used as fertilizer, together with
C. humeralis (Fabricius), which is piceous brown with a small yellowish-red spot
on the base of each elytron; this latter species is also widespread in South Africa.
Carpophilus dimidiatus is a reddish-brown species, often found in dried figs.
These are all small beetles varying in length from 2,3 to 3,5 mm.
A larger (about 4,8 mm long) and more robust dark-brown species, Aethina
castanescens (Fairmaire), was common in chicken manure in fowl pens in the
Cape Peninsula, together with larvae of house-flies, false stable-flies and lesser
house-flies, and the millepede Ommatoiulus moreleti (Lucas), but it was absent in
manure which harboured large populations of the lesser meal-worm Alphitobius
diaperinus (Panzer), and the skin-and-hide beetle Dermestes maculatus de Geer.
Families Cerambycidae and Curculionidae
Apodous larvae are found among long-horned and snout-beetles. In the
long-horned beetles (Cerambycidae) the larvae are elongate and somewhat
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
depressed. In the snout-beetles (Curculionidae), however, they are C-shaped
and usually have a hypognathous head. Larvae of long-horned beetles live in
wood and some are well-known pests. The adults, however, feed on flowers,
leaves, and bark or pollen of plants, and only one species belonging to the genus
Stenauxa was sometimes found among leaf litter and debris on the sand-dune
system of the coastal belt, together with the coppery oedemerid, Melananthis
senex Blair.
Adult snout-beetles, on the other hand, were common under cow-pats as
well as in leaf litter throughout the entire survey area, and were represented by at
least five subfamilies, of which the Eremninae predominated. At least three
well-known Eremnus species (Fig. 1D), viz. E. cerealis Marshall, E. setulosus
Boheman and E. atratus (Sparrman) (all endemic to the Cape Province according
to collection data) were found to be plentiful under dry pats, mainly in the
western Cape Province during the summer and autumn. Both E. cerealis
_ (speckled snout-beetle or grain-worm) and E. setulosus (grey snout-beetle) were
mostly observed under pats in fallow land and were collected only occasionally in
the veld. Eremnus atratus was also found at Hermanus during midwinter.
Eremnus pilosus Boheman, which resembles E. setulosus but is smaller, seems to
be restricted to the south-west of the Cape Province and was prevalent through-
out the spring. A small dark-brown Eremnus sp., about 4,5 mm long, was very
common near Saldanha Bay and Paternoster, but was never collected in any other
area.
The vegetable-weevil, Listroderes costirostris Schonherr, and Neocleonus
sannio (Herbst) (Fig. 1A) (both widely distributed in South Africa and overseas
(Csiki 1934, Voss 1973)), as well as Rhytirrhinus inaequalis (Fabricius) ((Fig. 1C),
which is endemic to the Cape Province, according to collection data), were mostly
found under dry cow-pats in the southern Karoo during the summer. However,
N. sannio was often found along the west coast under similar conditions, together
with Rhytirrhinus acerbus Boheman. During the late summer and early autumn
very large numbers of the vegetable-weevil, L. costirostris, may be present under
totally dry cow-pats in the sandveld area along the Cape west coast, particularly
during periods of drought. In certain areas up to three hundred specimens were
counted under a single large pat. Rhytirrhinus and Listroderes are more or less the
same size (7,5-7,8 mm long), but these two species of Rhytirrhinus are easily
distinguished by the strong carinae on the elytra. Neocleonus sannio is much
larger (12-13 mm), light grey in colour and marked with four to six brown
patches on the elytra.
A tiny, dull-black species of Ocladius, about 3,7 mm long and almost
globular in shape, with rugosoreticulate elytra and the pronotum with rows
of parallel, longitudinal, thin carinae connected with cross-ridges, was found
under dry pats in the Karoo during the late summer and autumn. At least one
species of Rhysoderes (Fig. 1B) was found to shelter under cow-pats near
Saldanha. It is 6,6—8,0 mm long, brownish grey, with head and pronotum rugoso-
punctate.
peli
SOUTH AFRICAN ARTHROPODS
lis.
inus inaequa
irrh
B. Rhysoderes sp. C. Rhyt
Neocleonus sannio.
A
Curculionidae.
Fig. 1.
D. Eremnus sp.
IAL ANNALS OF THE SOUTH AFRICAN MUSEUM
MORPHOLOGICAL DESCRIPTIONS AND BIOLOGICAL NOTES
The systematic arrangement followed is that of Britton (1970).
SERIES BOSTRYCHIFORMIA
Family Dermestidae
Small to medium-sized oval to somewhat elongate beetles, usually of sombre
coloration, often with coloured pattern due to the arrangement of scales on the
body. Head small and deflexed with clubbed antennae. Hind wings well de-
veloped, elytra covering the abdominal segments. Tarsal formula 5—5—5.
Of the nearly 500 species described, only about forty have been recorded
from southern Africa, of which at least ten species are of economic importance.
Hinton (1945) gives a good account (including illustrations) of the various species.
Dermestids are scavengers and feed on dried animal and vegetable matter with a
- high protein content, and some species are therefore attracted to decaying or-
ganic matter such as carcasses or cadavers, particularly during the formation of
butyric acid and other volatile fatty acids. Many species are found in nests of
insects and other higher animals, including birds and small mammals, often killing
the young, and a few are even predacious, feeding on the immature stages of
other insects such as the larvae of the wax-moth, Galleria mellonella Linnaeus, in
beehives.
Apart from the fact that the hairy larvae may cause dermatitis in sensitive
people, due to the liberation of histamine stimulated by urticating hairs (Smith
1973), there is also the danger of diseases such as anthrax (Bacillus anthracis
Koch) being spread by these beetles. The hairs of some species may remain in the
grain on which the larvae were feeding and may have serious effects when
swallowed (Morison 1925).
The larvae are easily recognized by their long body hairs, which may be
simple or modified, and members of the genus Dermestes have, in addition,
paired sclerotized urogomphi. They do extensive damage to stored foodstuffs
(grain and cereal products, dried fish, bacon, biltong, dried sausages, ham, etc.),
carpets, leather, feathers and hair, beeswax, clothes, skin and hides, bone,
insulating materials and museum specimens. Larvae and adults recovered from
Egyptian mummies had caused considerable damage to the embalmed bodies
(Strong 1981). At least eleven species of beetles were found in the skull of an
Egyptian mummy (Hope 1834), of which four were skin-and-hide beetles, viz.
Dermestes maculatus de Geer, D. peruvianus Castelnau, D. frischii Kugelann,
and D. roei Hope. Dermestid larvae will attack laminated boards or even
plaster-of-paris models if they contain glue of animal or vegetable origin, and
when ready for pupation they will perforate roof timbers, wooden panels, and
even lead sheeting. Under adverse conditions they may turn cannibalistic.
There are at least three vegetarian species of Trogoderma, of which the well-
known cosmopolitan Khapra beetle T. granarium Everts, is the most important.
It is a serious pest of stored grain elsewhere in the world, but is apparently not
SOUTH AFRICAN ARTHROPODS ZS
established in South Africa. However, it was reported from a brewery in Pieter-
maritzburg in malt imported from England as early as 1953, and from near
Pretoria in 1955 when a heavy infestation of this beetle and some other grain
insects occurred in a store in sweepings that had been allowed to accumulate for
about three years.
The carpet-beetles include at least six cosmopolitan or nearly cosmopolitan
species that have been recorded from South Africa, including the widely distrib-
uted varied carpet-beetle, Anthrenus verbasci (Linnaeus), also known as the
museum-beetle, which is so destructive to dried museum specimens, and the true
museum-beetle, A. museorum (Linnaeus), reported in the past only from South
West Africa. The furniture carpet-beetle, A. vorax Waterhouse, which is very
similar to A. verbasci, but is easily separated from the latter by the inner or
mesal margin of the eyes, which is excised or interrupted in the middle (entire
in A. verbasci), has been observed in nests of the social weaver-bird (Phile-
tairus socius (Latham)). The larvae feed on feathers and other debris. One of
the more common species found in carpets, bird carcasses and poultry manure in
the western Cape Province is the cosmopolitan black carpet-beetle, Attagenus
piceus (Olivier), which is more elongate than species of Anthrenus; it is occasion-
ally found in association with the indigenous Attagenus jucundus Peringuey
and the nearly cosmopolitan A. gloriosae (Fabricius) in carpets in the Cape
Peninsula.
Skin-and-hide beetles are common in decaying carcasses, both on the
beach and inland, in fish-meal, dried fruits, animal droppings, and even horn,
feathers, and biscuits. Four cosmopolitan species are recorded from South Africa,
of which Dermestes maculatus and D. peruvianus are the most numerous in the
western Cape. Dermestes ater de Geer, although widespread, is a more northern
species, whereas the larder-beetle, D. lardarius Linnaeus, is only found occasion-
ally in the Cape Town docks, mostly in dried foodstuffs such as macaroni and
walnuts.
Dermestes maculatus de Geer
DESCRIPTION
Adult
Previously described by Hinton (1945). Blackish, dorsally clothed with de-
cumbent fulvous to greyish hairs, with mostly head and a broad band along each
lateral margin of pronotum covered with longish white hairs. Apex of each
elytron serrate and sutural angle produced in form of a strong tooth, absent in the
other two species described here. Lateral impressed line of first sternite curved
inwards near base. Length of specimens examined varying from 5,5 to 10 mm.
A native of Europe and widely spread in most parts of the world (Hinton
1945). It is the most common skin-beetle around Cape Town and is also well
established on most of the islands along the South African south and west coasts,
including Dyer, Bird, Marcus, Jutten, Ichaboe, and Malgas islands.
214 ANNALS OF THE SOUTH AFRICAN MUSEUM
Larva (Fig. 2A)
General descriptions are given by Rees (1943), Hinton (1945), and Peterson
(1967). Body form of Dermestes maculatus larva elongate, semi-circular; tergites
sclerotized and piceous to dark brown with broad, pale yellowish-white median,
longitudinal line over dorsum, representing ecdysial suture. Head hypognathous.
Tergal plates covered with medium-long to very long spinulate setae, some hairs
on third to eighth abdominal segments somewhat longer than others. Spinulate
hairs on ventral side paler and shorter. Integument smooth, without spinules
except for short branched (spinulate) spines on certain areas. A transverse row of
strong pointed tubercles present on anterior margin of dorsal sclerotized plates of
abdominal segments 4—9 (Fig. SA—B). Spiracles well developed, mesothoracic
one the largest, oval (Fig. 2C); first three abdominal spiracles less oval and
situated on a non-sclerotized membrane below sclerotized tergal plates, those on
fourth to eighth segment smaller than other abdominal spiracles, almost circular
- and situated on the sclerotized tergal plates (Fig. 5A); in pale-coloured larvae
they are often less sclerotized than others (Fig. 2D). Legs pale brownish to
brown, tarsungulus (Fig. 3L) dark brown, and with short narrow keel-like projec-
tion ventrally on each side of which is a long fine seta. Setation of posterior face of
tibia as in Figure 3L; characteristic of D. maculatus is the long fine posterodorsal
preapical seta on tibiae. Length of full-grown larva about 15 mm.
Head (Fig. 2E—G)
About as wide as long, or only slightly wider than long, sides and hind margin
dorsally fairly convex. Frontal and epicranial (coronal) sutures distinct; frons
strongly sclerotized except for narrow strip along anterior border and on each side
in front of antennae; each side of frons with strong conical tubercle. Epicranium
also strongly sclerotized, except for small area around ocelli; latter six in number,
on each side arranged in two almost vertical parallel rows. Setation more or less
as illustrated. Antennae (Fig. 2B) three-segmented with whitish cone-shaped
basal articulating membrane; first segment slightly more than half the length of
second, wider than others; with about six setae situated dorsally some distance
from its apex; second segment with sensory appendix (sensory cone) at apex and
an oval sensory spot near middle. Third segment small, about one-third the length
of second, with sensory pegs at apex.
Labrum and clypeus
Labrum slightly more than half the length of clypeus, narrower than latter,
its anterior margin fairly deeply and angularly excised in middle. Clypeus
trapezoidal, much narrower in front than behind; anterior margin slightly con-
cave, preclypeus without setae, except for medium-long seta on each side near
lateral margin; postclypeus with about six to eight medium-long setae.
Epipharynx (Fig. 2H) with proximal sensory area composed of about six
large and ten small sensory papillae arranged in two almost parallel rows.
Epipharyngeal scleromes parallel, tapering and somewhat diverging posteriorly.
SOUTH AFRICAN ARTHROPODS 215
=
Y, |
Fig. 2. Dermestidae. Dermestes maculatus larva. A. Full-grown, left lateral view. B. Antenna.
C. Mesothoracic spiracle. D. Abdominal spiracles, segments 1-3, 4, 8. E. Head, dorsal view.
F. Head, ventral view. G. Head, left lateral view. H. Epipharynx. I. Left mandible, dorsal
view. J. Right mandible, dorsal view. K. Left maxilla, dorsal view. L. Labium, ventral view.
M. Labium, dorsal view. N. Labium, left lateral view. O. Left maxilla, ventral view.
216 ANNALS OF THE SOUTH AFRICAN MUSEUM
Broad median lobe setae about five in number on each side, lateral lobe setae
longer and finer and covering each lateral margin to a point well beyond its
middle. Also two sensory pegs situated in median V-shaped excision below two
dorsal median setae.
Mandibles (Fig. 2I-J)
Broad, almost triangular and slightly less than one-third the length of head.
Cutting edge tri-dentate, apical tooth (ti) somewhat larger than others. Molar
area absent, but represented by large stiff process of retinaculum and brush of
hairs or penicillus. Lateral face with demi-scrobe with about four setae of which
the distal two are longer than others; also transverse row of hairs near base of
mandible on lateral face, with median hairs the longest.
Maxillae (Fig. 2K, O)
Galea and lacinia united, only free at extreme apex, the boundary lines
clearly indicated on both sides. Lacinia with forked spur at apex and dorsally also
with brush of fine fulvous hairs, otherwise devoid of setae. Galea lobe-like with
brush of fine fulvous hairs apically and four to five short setae on external lateral
margin just below brush of hairs, as well as one somewhat longer seta on ventral
side near palpifer. Stipes broad, almost quadrate with some setae on ventral side
of which one is much longer than rest; dorsally without setae; also some
medium-long setae along its ridge-like exterolateral margin. A narrow juxtastipes
present, demarcated by sclerotized margin of stipes. Palpifer without setae.
Maxillary palp four-segmented, the segments somewhat tapering towards apex;
first and second segments of about equal length and somewhat shorter than third
and fourth in specimens examined; first segment with a single ventral seta, the
penultimate one with two setae, one lateral and one ventral; apical segment with
some sensory pegs at apex. Subcardo and precardo (alacardo) clearly indicated by
sclerotized demarcation line; former with only about three, latter with about
seven setae.
Labium (Fig. 2L—N)
Submentum and gula fused, mentum almost rectangular, longer than wide
with setal pattern as indicated in figure, including longitudinal row of setae on
lateral face (paramentum). Ligula cleft in middle, ventrally with about five setae
on each lobe as indicated; hypopharyngeal sclerome as a curved sclerite and
raised above the palpi in all specimens examined, superlinguae beset with minute
setae. Labial palpi two-segmented, tapering towards apex, segments of about
equal length, without setae except for sensory pegs on apical one.
Pupa (Fig. 3A-B, E-G, J)
Somewhat elongate, about two and a half times longer than wide when seen
from above, creamy white in colour; completely covered with fine medium-long
fulvous hairs, each hair growing from a small rounded tubercle, which is coloured
SOUTH AFRICAN ARTHROPODS 2h
¢
G 0 ae ree mo vy
Fig. 3. Dermestidae. A. Dermestes maculatus pupa, left lateral view. B. Dermestes maculatus
pupa, pronotum, dorsal view. C. Dermestes ater pupa, pronotum, dorsal view. D. Dermestes
peruvianus pupa, pronotum, dorsal view. E. Dermestes maculatus, antenna of pupa, left lateral
view. F. Dermestes maculatus, abdominal apex of pupa showing developing genital capsule,
ventral view. G. Dermestes maculatus, head of pupa, dorsal view. H. Dermestes peruvianus,
urogomphus of pupa, left lateral view. I. Dermestes ater, urogomphus of pupa, left lateral view.
J. Dermestes maculatus, urogomphus of pupa, left lateral view. K. Dermestes peruvianus, head
of pupa, dorsal view. L. Dermestes maculatus, tarsungulus and tibia of left front leg of larva,
posterior view. M. Dermestes ater, tarsungulus and tibia of left front leg of larva, posterior view.
N. Dermestes peruvianus, tarsungulus and tibia of left front leg of larva, posterior view.
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
yellowish red; tubercles and their coloration more prominent on median dorsal
side of pupa, particularly in newly formed pupae; those on pronotal disc some-
what larger than others. The five so-called ‘gin-traps’ on third to seventh abdomi-
nal terga coloured reddish and very distinct, even in old pupae. Spiracles large,
almost circular on abdominal segments 1-6, that on seventh smaller and less
obvious. Two urogomphi vertical and almost parallel. Developing genital capsule
in form of two swellings as indicated in figure.
Characteristic of the pupae of all three species of skin-and-hide beetles
examined are the six to ten setae on the second ringlike segment of the antennal
sheath (Fig. 3E). These setae are also present in the pupae of Attagenus piceus, in
which case there are six ‘gin-traps’. In all the pupae of Anthrenus verbasci that
were examined these setae and the ‘gin-traps’ were absent.
BIOLOGY
In most species of dermestids the adult beetles need food and access to a
water supply for satisfactory reproduction. This was also found by Dick (1937),
who states that if beetles were allowed to drink they would continue to lay eggs
for 72 days. This is not the case with the smaller carpet-beetles such as Anthrenus
verbasci, which are able to complete their adult life and lay eggs without any
feeding.
The eggs of Dermestes maculatus found in decaying carcasses were dirty
white and minutely and longitudinally striate and also reticulate when viewed at a
certain angle. Most eggs examined varied from 1,30 by 0,54 to 2,04 by 0,92 mm
and were fairly dull.
In larvae reared in the laboratory on their natural food supply, the larval
life-span varied from 31—42 days during January to March, the pupal stage being
7-11 days. During April to May they needed about 50 days to mature and the
pupal stage was about 35 days. Pupation occurs in the larval skin in the soil
without the formation of an earthen cell.
The newly emerged beetle is pale yellowish white with the abdominal apex
reddish; pronotum pale reddish surrounded by a yellowish-white, broad border.
Legs pale, trochanters, tarsi and articulation points of femur, and tibiae reddish;
head also pale reddish, eyes brown, antenna reddish with the apical three seg-
ments piceous brown, and apices of mandibles black. After a few hours they
assume their normal dark coloration.
As indicated previously, during the act of pupation the larvae will perforate
any suitable material for this purpose and may cause severe damage to timber in
buildings. In the past various instances have been recorded where larvae of this
species caused damage amounting to thousands of rands where laminated boards
and wooden panels contained animal glue. It is also possible that the larvae may
consume some of the wood as part of their diet. This fact is also stressed by
Bedwell (1931) when he refers to the smooth hollowed-out parts of the timber
and the presence of the larvae in various stages of development.
During the surveys it was observed that the larvae of D. maculatus very often
SOUTH AFRICAN ARTHROPODS 219
feed on their own pupae and they in turn are heavily preyed upon by larvae of the
ham-beetles, particularly Necrobia rufipes, which sometimes occur together with
this species.
Dermestes ater de Geer
DESCRIPTION
Adult
Previously described by Hinton (1945). Dark castaneous to piceous, integu-
ment somewhat more shiny than in Dermestes maculatus; body hairs fulvous.
Lateral impressed line on first sternum widely curved inwards, only parallel to
lateral margin for about half its length (in D. maculatus it is parallel for most of its
length). Elytral apices without serrations and without sutural teeth. Length
varying from 8,3 to 9,5 mm.
Indigenous to America (Fauvel 1889) and widely spread in the world (Hinton
1945). Only occasionally observed in the Cape Town docks.
Larva
Very similar to that of D. maculatus (see Fig. 2) in size, body form and
setation including spiracles, except for pro-mesothoracic spiracle (Fig. 4B), which
in D. ater bears about twelve almost equally developed ventral setae, whereas in
all specimens of D. maculatus that were examined, there were only one strong
and about five weaker setae. In this species, as in D. maculatus, the first three
abdominal spiracles are situated on a non-sclerotized membrane below sclero-
tized tergal plates, but the abdominal segments are devoid of a transverse row of
strong tubercles (Fig. 5C-D). Urogomphi straight (Fig. 5D—in D. maculatus
slightly curved upwards and anteriorly), and tibiae (Fig. 3M) each with stout
dorsal spine on posterior face near apex (in D. maculatus it is replaced by long
slender seta); short spine-like ventral seta present on each side of base of tarsun-
gulus (in D. maculatus these setae are long). In most specimens examined the
median, longitudinal yellowish band over the dorsum in the form of a thin line,
forming a diamond-shaped patch in middle of each segment.
Head (Fig. 4A)
Very similar to that of D. maculatus in shape, sclerotization and setation, but
lacking tubercles on frontal area. Six ocelli on each side arranged in more or less
two groups of three each, as illustrated. Frontal and coronal sutures distinct.
Head of most specimens examined with somewhat more setae than in D. macula-
tus (see Fig. 2E—G), particularly on frontal area. Antennae also very similar to
those of latter species, including six dorsal setae on first segment; the sensory
appendix, however, seems to be somewhat smaller.
Labrum and clypeus
Very similar to those of D. maculatus, including emargination in middle
of anterior margin of labrum. Epipharynx (Fig. 4E) also similar and with double
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. Dermestidae. A-J. Dermestes ater larva. A. Head, left lateral view. B. Pro-
mesothoracic spiracle, left lateral view. C. Left mandible, dorsal view. D. Right mandible,
dorsal view. E. Epipharynx. F. Left maxilla, dorsal view. G. Labium, ventral view.
H. Labium, dorsal view. I. Left maxilla, ventral view. J. Labium, left lateral view.
K-S. Dermestes peruvianus larva. K. Head, left lateral view. L. Left mandible, dorsal view.
M. Right mandible, dorsal view. N. Left maxilla, dorsal view. O. Labium, ventral view.
P. Labium, dorsal view. Q. Left maxilla, ventral view. R. Epipharynx. S. Labium, left
lateral view.
SOUTH AFRICAN ARTHROPODS DIA
row of proximal sensory papillae, the front six large, arranged in two groups of
three each in most specimens seen; posterior row with about fourteen minute
papillae. Epipharyngeal scleromes also similar to those of D. maculatus (see
Fig. 2H); broad median lobe setae about five in number on each side; lateral lobe
setae spear-shaped and covering each lateral margin to a point just beyond its
middle.
Mandibles (Fig. 4C-D)
Very similar to those of D. maculatus, including demi-scrobe and transverse
row of hairs; however, retinaculum much narrower and penicilli much broader in
most specimens examined than in that species.
Maxillae (Fig. 4F, I)
Similar to those of D. maculatus (see Fig. 2K) and without any setae on
dorsal side, except for hair brushes on galea and lacinia. Setal pattern of ventral
side similar to that of latter species. Demarcation line between galea and lacinia
fairly distinct, the spur as in D. maculatus. Maxillary palp with third and fourth
segments of about equal length and both slightly longer than first and second;
apical segment narrowest; setal pattern, subcardo and precardo exactly as in
D. maculatus, except no setae could be traced on subcardo.
Labium (Fig. 4G—-H, J)
Very similar to that of D. maculatus, including shape, setal pattern, labial
palpi and hypopharynx.
Pupa
Also very similar to that of D. maculatus in colour, body form, size and shape
of developing genital capsule, but differing from it by the somewhat smaller hair
tubercles, which are more or less confined to median area of tergites, particularly
the pronotum (Fig. 3C), even in older pupae. In D. ater the urogomphi (Fig. 31)
are also more curved anteriorly and are not straight. In most specimens examined
there seem to be slightly more setae on the body, and the setae are somewhat
finer and more slender.
Dermestes peruvianus Castelnau
DESCRIPTION
Adult
Previously described by Hinton (1945). Dark castaneous brown, clothed with
short decumbent fulvous hairs. Integument somewhat more shiny than in Der-
mestes maculatus. Lateral impressed line on first sternite straight and parallel to
lateral margin. Elytral apices entire, not serrate and also lacking sutural teeth.
Length 6,6-11,0 mm.
D2? ANNALS OF THE SOUTH AFRICAN MUSEUM
Probably a native of South America and widely distributed through the
major countries of the world (Hinton 1945). During surveys it was not found on
the islands along the Cape coasts; however, it is almost as common as D. macula-
tus in carcasses and fish-meal accumulations around Cape Town.
Larva
Very similar to that of D. maculatus (see Fig. 2) in size, body form and
setation, including the form of the spiracles; D. peruvianus, however, has pro-
mesothoracic spiracle with more setae (about twelve) of equal size as in D. ater,
and only first two abdominal spiracles are situated on a non-sclerotized mem-
brane below sclerotized plates of the two segments (Fig. SE); also transverse row
of strong tubercles present on anterior border of sclerotized tergal plates on
peas i
Fig. 5. Dermestidae. A. Dermestes maculatus \arva, left lateral view of spiracles and tergal
plates on segments 3 and 4. B. Dermestes maculatus larva, left lateral view of segments 9 and 10.
C. Dermestes ater larva, left lateral view of spiracle and tergal plate on segment 4. D. Dermestes
ater larva, left lateral view of segments 9 and 10. E. Dermestes peruvianus larva, left lateral view
of spiracles and tergal plates of segments 2 and 3. F. Dermestes peruvianus larva, left lateral view
of segments 9 and 10.
abdominal segments 3 to 10 (Fig. SE-F). Urogomphi in this species (Fig. 5F)
fairly straight, their apices curved caudally and downwards and, when seen from
above, slightly divergent as in D. ater. Tarsungulus (Fig. 3N) with broad base, on
each side with long slender ventral seta, tibiae with long slender posterodorsal
preapical seta as in D. maculatus.
SOUTH AFRICAN ARTHROPODS DDG
Head (Fig. 4K)
Very similar to that of D. maculatus (see Fig. 2E—F) in form, sclerotization
and setation, including two tubercles on frontal area and distinct frontal and
coronal sutures. Six ocelli on each side appearing somewhat smaller and arranged
in two groups of three each. Antennae also similar to those of latter species,
including sensory appendix, but in most specimens examined oval sensory spot
placed on posteroventral side of second segment (in D. maculatus it is more
ventral in position).
Labrum and clypeus
Very similar to those of D. maculatus including setation and labral emargina-
tion. Epipharynx (Fig. 4R) as in D. maculatus (see Fig. 2H) with proximal
sensory area consisting of an anterior row of six large and a posterior row of about
twelve minute papillae. Median setae about three to four in number in most speci-
mens seen and rather broad, their apices either truncate or rounded, the setae
somewhat broader than in D. ater and rather similar to those of D. maculatus.
Mandibles (Fig. 4L—M)
Very similar to those of D. maculatus, including broad retinaculum, rather
narrow penicillus and transverse row of hairs near mandibular base; each demi-
scrobe also with about four shorter setae. Apical tooth (ti) the largest.
Maxillae (Fig. 4N, Q)
Similar to those of both D. maculatus and D. ater in form and setation,
including maxillary palpi, forked spur on lacinia and hair brushes on both lacinia
and galea. In this species there are also about three setae on subcardo.
Labium (Fig. 40-P, S)
Very similar to that of D. maculatus including setal pattern and hypo-
pharynx. Labial palpi also similar, but differ from those of both D. maculatus and
D. ater by the first segment which bears at least two ventral setae.
Pupa
Similar to D. maculatus but the hair-bearing tubercles, particularly on the
pronotum (Fig. 3D), although of about the same size, are more or less confined to
the median area in D. peruvianus, and in the few specimens examined the hind
margin of the mesonotum is more rounded in middle (in D. maculatus it is more
pointed or rather acute). Urogomphi (Fig. 3H) more curved anteriorly as in
D. ater and not straight as in D. maculatus.
SERIES ELATERIFORMIA
Family Cantharidae
Small to medium-sized, elongate, somewhat flattened leather-winged
beetles, usually with reddish and bluish coloration. Head generally narrower than
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
pronotum and elytra, deflexed and with filiform antennae. Elytra covering the
abdominal segments; tarsal formula 5—S—5. Luminous organs absent. According
to Arnett (1963) they are in some ways the most primitive of beetles.
About fifty species have been recorded from southern Africa; their habits are
largely unknown. Only one species of the genus Afronycha has been observed in
animal dung during the surveys.
Afronycha picta (Wiedemann)
DESCRIPTION
Adult (Fig. 6A)
Dark olive green with orange-red head and pronotum marked with dark
green. Dull, covered all over with fine, short greyish hairs. Legs orange red.
Antennae usually dark; first two segments pale orange red. Length 9,1-11,6 mm.
Wittmer (1960) gives the distribution as Cape Town, Rondebosch, Camps
Bay and Paarl. Delkeskamp (1977) mentions only Natal. According to collection
data, however, it is widely distributed in the Cape Province.
Larva (Fig. 6B)
A general description of a cantharid larva is given by Peterson (1967). Full-
grown larva of Afronycha picta measures 17-18 mm long and is velvety brown
with somewhat bluish tinge on thoracic segments; each body segment with pale
yellowish-brown marks and stripes and thin pale line on lateral margin. Tenth
abdominal segment with small fleshy protuberance (Fig. 6L) acting as a proleg.
Thoracic legs (Fig. 6J) well developed; claws (Fig. 61) simple, long and acute,
with two small spines near base on posterior side and one on anterior side. Each
body segment with gland opening on each side. Spiracles (Fig. 6K) similar to that
illustrated by Boving & Craighead (1931), opening almost circular with black,
triangular area anteriorly, which, according to above authors, possibly represents
the reduced air-tubes. Spiracular opening furnished with fine hairs. Cantharid
larvae are campodeiform and orthosomatic.
Head (Fig. 6E)
Brownish with one well-developed ocellus on each side behind antenna.
Subnasal sclerite sinuate on each side, with small tooth in middle. Epicranial
halves fused ventrally. Antennae three-segmented; second segment about twice
as long as first; third very small; second segment with a sensory appendix
(Fig. 6F). Mandibles (Fig. 6G) almost falcate, nearly twice as long as wide, with
two acute apical teeth; each mandible with a dorsal longitudinal row of fine hairs
extending from preartis; lateral margin fairly strongly carinate; ventral side of
teeth with conduit for juice. Mala (Fig. 6F, H) small and with long terminal hair.
Mentum and stipes (Fig. 6F) clearly marked and fairly well sclerotized; premen-
tum almost triangular. Maxillary palpi appearing four-segmented, but according
to Peterson (1967) only three segments are present; apical segment about half as
SOUTH AFRICAN ARTHROPODS D225
Fig. 6. Cantharidae. Afronycha picta. A. Adult. B. Larva, dorsal view. C. Pupa, ventral view.
D. Detail of last segment of pupa, ventral view. E-—L. Larva. E. Head, dorsal view. F. Head,
ventral view showing mentum and stipes. G. Mandibles, dorsal view. H. Hypopharyngeal area,
dorsal view to show straining hairs. I. Claw, enlarged to show spines. J. Right mesothoracic leg.
K. Right abdominal spiracle to show triangular area and hairs. L. Apical abdominal segments,
left lateral view.
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
wide as preapical one and tapering towards apex; the latter segment the shortest
and ring-like. Hypopharynx (Fig. 6H) with at least three longitudinal rows of
straining hairs.
Pupa (Fig. 6C)
Whitish yellow with reddish eye-spots and extreme apices of mandibles
reddish. Specimens examined measure 10,1 mm long and 4,6 mm broad across
widest part. Extreme apex of abdomen with two short spines or dents. Develop-
ing genital capsule (Fig. 6D) visible as large convex or rounded tubercle. As
imago develops pupa becomes darker and apices of mandibles turn almost black;
abdomen remains yellowish white and wing covers have piceous tinge.
BIOLOGY
Many species of cantharids are predacious, some feeding on aphids, others
-on the eggs of grasshoppers and on the larvae of Lepidoptera and Coleoptera;
others again are phytophagous; some are omnivorous. The adult beetles often
visit flowers where they feed on pollen. The larvae are free-living and predacious,
preying on the larvae of other insects.
Young larvae of Afronycha picta were collected along the west coast of the
Cape Province in fairly fresh as well as semi-fresh cow-pats during midwinter.
These larvae are greenish in colour, dotted with tiny black and larger dark-grey
spots. As they become older, the colour turns darker, until they assume their
normal brownish-velvet appearance. Mature larvae were found in pats during
August. They are fairly sluggish and feed on the coprophagous insects, including
larvae of dung-flies (Orthellia spp.) and flesh-flies (Sarcophaga spp.). They have
the habit of rolling around with the struggling prey until movement of the latter
subsides, by which time half of the prey may have been devoured. The larvae
were seen feeding on the juices oozing from wet dung.
The development of the larvae seems to be fairly slow and pupae were only
recovered at the end of August and beginning of September from larvae collected
in July. The pupal stages lasted about 11 days in the laboratory (19-22 °C); the
beetles then emerged during September. In the Darling district they were also
observed to emerge from cow-pats during May.
Family Melyridae
Small to medium-sized beetles, often with brilliant blue or red coloration, the
head nearly as broad as pronotum and with serrate antennae. Elytra entire and
tarsal formula 5—5—5 (rarely 4—5—5).
The Melyridae were previously regarded as a subfamily of the Dasytidae but
have now been elevated to include the Malachiidae as well as the Dasytidae, the
latter containing the most primitive living members of this group. Some author-
ities, such as Crowson (1952, 1964), regard the true position of this family to be in
the superfamily Cleroidea and consider its resemblance to the family Cantharidae
merely as a consequence of convergence. Melyrids most closely resemble can-
SOUTH AFRICAN ARTHROPODS D2F,
tharids both in structure and habits; however, the structure of the aedeagus of its
members bears a clear resemblance to that of the superfamily Cucujoidea.
Melyrid larvae on the other hand seem to be more closely related to the super-
family Cleroidea if certain characters such as the presence of a gular area,
urogomphi and abdominal glands (also present in cantharids) are considered.
However, the larvae are more free living and are either scavengers or feed on
other insects, their larvae and eggs.
The adults are most probably carnivorous, although they are often found on
flowers, where some feed on pollen. A few species are phytophagous such as the
spotted maize-beetle, Astylus atromaculatus (Blanchard), a native of South
America which is widespread in the Republic of South Africa. The adults of
A. atromaculatus attack the flowers of maize, sorghum, etc., and are gregarious,
accumulating in large numbers on the plants; it is also poisonous to cattle and
sheep (Bedford et al. 1974).
A small metallic-blue and orange-coloured species, Colpometopus basicornis
(Fairmaire), is common in stranded kelp and was also found to feed on the tissues
of decaying animal carcasses on the beach, together with individuals of the larger
Melyris viridis.
Melyris viridis Fabricius
DESCRIPTION
Adult (Fig. 7A)
Dull metallic green or sometimes with elytra coppery green. Each elytron
with three distinct striae; whole insect dorsally covered with piliferous punctures.
Hairs black and short. Pronotal disc on each side with longitudinal carina extend-
ing Over anterior two-thirds and ending in raised frontal margin. Pronotum also
with shallow, median, longitudinal impression, which is not very obvious in small
specimens. Legs setose, lateral side of body fringed with short, fine, black hairs.
Pronotum narrower than elytra. Length 9,6-13 mm.
Widely spread along the western and north-western coasts of the Cape
Province.
Larva (Fig. 7B)
General description of a melyrid larva is given by Peterson (1967).
Melyris viridis larva subcylindrical, with nine visible abdominal terga; head
small. About 19 mm long when fully grown and light brownish red in colour;
vistiture consists of long, golden hairs. Prothoracic segment with broad, brown to
blackish sclerotized shield, divided by thin, pale reddish, longitudinal (ecdysial)
line. Meso- and metathoracic segments each with lateral brownish sclerotized
area. Head, legs, and ninth abdominal tergum blackish; latter sclerotized dorsally
and bearing two urogomphi which are curved upwards. Urogomphus with small
tooth laterally, some distance from apex (Fig. 7G). Legs well developed, increas-
ing in size from front to back; coxa, femur, and tibia well demarcated and
furnished with long hairs; tarsungulus (Fig. 7F) moderately long, acute, and with
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 7. Melyridae. Melyris viridis. A. Adult. B-G. Larva. B. Dorsal view. C. Head, dorsal
view. D. Head, left lateral view to show ocelli. E. Mouth-parts, ventral view. F. Right
mesothoracic leg. G. Eighth and ninth abdominal segments, left lateral view.
H. Pupa, ventral view. I-N. Larva. I. First abdominal segment to show the extruded defensive
gland. J. Antenna. K. Maxillae and labium, dorsal view. L. Mandibles, dorsal view.
M. Anterior part of labrum, dorsal view. N. Apex of lacinia showing tooth (highly magnified).
SOUTH AFRICAN ARTHROPODS 229
single long ventral seta on enlarged base. Two pairs of protrusible, lateral,
defensive glands present (Fig. 71); one pair on first and other pair on eighth
abdominal segment; each gland normally indicated by narrow sclerotized slit just
behind the spiracle (Fig. 7G). When larva is irritated, glands are evaginated.
Crowson (1964) describes the same number of glands in Astylus atromaculatus
and in Melyris rufiventris Boheman. Spiracles annular (Fig. 71), small, and in
most of the mature specimens examined opening is only 0,08 mm in diameter; the
mesothoracic spiracle somewhat larger, measuring about 0,10 mm. Anus present
as transverse slit, surrounded by single upper and double lower lip (divided by
transverse furrow into two parts).
Head (Fig. 7C-E)
Slightly wider than long, black; epicranial suture Y-shaped and epicranial
stem (coronal suture) well developed. Both frons and epicranium beset with long
golden hairs. Four ocelli on each side, with posterior one largest. Antennae
(Fig. 7J) three-segmented, first two segments somewhat compressed laterally and
retractable into antennal base; third segment small, with single medium-long seta
and some sensory pegs at apex. Second antennal segment with triangular raised
area ventrally near apex; also with three medium-long apical setae of which two
are lateral and one dorsal in position. Gula clearly demarcated and about as wide
as or slightly wider than labium.
Small larvae, only 6,0 mm long, showed same characteristics as mature ones,
except for smaller number of setae on labrum.
Labrum and clypeus
Both well developed; labrum about as long as clypeus, and almost semi-
circular in specimens seen. It bears median transverse row of long setae as well as
second transverse row of shorter setae near extreme anterior margin, each seta
placed on a small black sclerotized area (Fig. 7M). Ventral side of labrum with six
strong palus-like setae close to anterior margin (clearly seen in Fig. 7M). In most
Specimens seen, epipharynx bears six large papillae, medially arranged in two
rows; four in anterior and two in posterior one; also two small papillae posterior
to latter row. Clypeus with only single long seta on each side.
Mandibles (Fig. 7L)
Slightly longer than wide; fairly broad and robust in dorsal view. Dorsal
carinae almost absent, mandibles rounded dorsoventrally in this area; however,
very weak oblique scrobis present with eight or ten setae, of which proximal one
is longest. Stridulatory areas absent; prostheca well developed and situated near
the base. True molar area absent, cutting edge broad and almost quadridentate in
most specimens. Fairly acute apical (ti and ts) and preapical teeth (tz and ts) are
present, latter situated ventrally; proximal teeth (ts and ts) smaller, dorsally
situated, usually slightly bidentate and apparently functioning as molar area.
Portion of cutting edge between ventral and dorsal teeth shallowly excavated.
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxillae (Fig. 7E)
Short, galea and lacinia united and almost inseparable. According to Béving
& Craighead (1931) lacinia has no spurs in Melyridae. However, apex of lacinia is
angular with minute tooth connected to internal tube (Fig. 7N), visible only under
high magnification. Galea globular, with large number of fine setae on dorsal
side, but ventrally devoid of hairs; two long ventral setae, however, present just
posterior to palpal bases. No unci observed. Maxillary palp three-segmented;
second segment with lateral and third segment with dorsal seta as well as some
sensory pegs. Cardo transverse and with single long seta. Stipes long and clearly
demarcated ventrally, with about six long setae apically and two basal setae;
dorsally devoid of setae.
Labium (Fig. 7E, K)
Submentum and mentum (apparently the first and second prementum of
~ Anderson (1936)) fairly distinct; the former with three to five long setae on each
side, the latter with two to three setae. Prementum of labium visible as two
sclerites, each with a single seta; labial palpi two-segmented; gula convex. Glossa
(or ligula) pointed, with four sensory spots. Basally two swollen lobes may
represent superlinguae of hypopharynx.
Pupa (Fig. 7H)
Dark reddish with whitish antennae, legs, and elytra. Whole body abun-
dantly covered with long golden-brown hairs, particularly on dorsal side. Body
has bluish violaceous shine and after few days two brownish eyes become visible.
The developing genital capsule present as two short conical projections; extreme
apex also with two short, fleshy protuberances. Length 10,5—12,9 mm.
BIOLOGY
Adult beetles of Melyris viridis were collected on the beaches and foredune
system along the west coast of the Cape Province from Kommetjie to Elands Bay,
almost throughout the year. They feed on the pollen of flowers of Arctotheca
populifolia and Tetragonia decumbens and were seen to copulate in large numbers
during midwinter (June) and also during the early summer (November) when
peak emergence occurred during the surveys. They are attracted to decaying
carcasses and cow-dung and were often observed to nibble holes in the leaves and
flowers of the above-mentioned plants.
Fully developed eggs dissected from adults in August were brownish red,
oval, and varied in size from 0,8 by 0,44 mm to 1,42 by 0,64 mm; they were
almost shiny and very superficially reticulate. The eggs are apparently laid during
the winter, the mature larvae burrowing into the sand from September to Novem-
ber; eggs that were laid during the summer gave rise to full-grown larvae during
the winter. Some of the larvae kept under observation passed the summer in
sheltered spots in the sand or under dry kelp, and even under dry cow-pats, and
pupated the following April, the beetles then emerging in June. These larvae are
SOUTH AFRICAN ARTHROPODS 23K
therefore present on the beach for the greater part of the year, except possibly for
a short period during the winter and again during the summer. Observations
made during 1976 indicated that there were two generations per year with
overlapping of adults and larvae. Pupation took place 50-100 mm deep in the
sand under plants growing on the upper boundary of the beaches. During mid-
summer full-grown larvae and beetles can often be found about 15 cm deep in the
sand below plants growing on the beach.
The larvae are omnivorous and feed on semi-dry parts of decaying kelp,
partly dry cow-dung, decaying red-bait (Pyura stolonifera (Heller)), bird and
other animal carcasses, as well as on leaves and flowers of such plants as Arcto-
theca populifolia and certain vygies, and even on vegetables, such as pumpkins,
thrown on the beaches. When resting in the sand they were seen to feed on the
beach-fleas, Talorchestia australis Barnard and T. capensis Dana.
SERIES CUCUJIFORMIA
Family Tenebrionidae
Small to fairly large, almost round to elongate beetles, varying from white to
brown or black, sometimes even with reddish marks. The prognathous head is
narrower than pronotum, elytra usually covering the abdomen and often with
striae. Tarsal formula 5—5—4.
According to Watt (1974) the primitive forms are relatively large and both
larval and adult life is usually long. It is one of the largest families of the
Coleoptera, the largest forms occurring in the genus Psammodes, measuring up to
60 mm in length. Most of them are adapted to resist desiccation and they are
therefore well represented in arid or semi-arid areas and may occur almost
anywhere, including birds’ nests; however, they are apparently not adapted to
living in water.
The adults generally feed on dead vegetable and sometimes also on animal
matter. The larvae of most species, however, will feed on living plant tissues—
particularly under adverse conditions. When crowding occurs the larvae of most
species become cannibalistic. Wings are well developed in the smaller species,
particularly the stored-food pests such as the lesser meal-worm, Alphitobius
diaperinus (Panzer), the common meal-worm, Tenebrio molitor Linnaeus, the
horned flour-beetles, Gnathocerus spp., the confused and rust-red flour-beetles,
Tribolium confusum du Val and T. castaneum (Herbst), the dark flour-beetle
T. destructor Uyttenboogaart, the long-headed flour-beetle Latheticus oryzae
(Waterhouse) as well as in the dusty surface-beetles Gonocephalum spp. The
psammophilous members and toktokkies (Psammodes spp.) on the other hand
are apterous. Some tenebrionid beetles are able to protect themselves by secre-
tions from defensive glands.
Psammodes species, particularly P. granulatus Solier, were common on the
sand-dune system together with Zophosis acuta Wiedemann. The latter is an
endemic southern Cape dune species, which was also occasionally attracted to
232 ANNALS OF THE SOUTH AFRICAN MUSEUM
decaying kelp on the beach, most probably because of the high water content of
the seaweeds, as these insects seem to favour damp substances; Z. acuta often
appeared under semi-dry cow-dung some distance from the shore.
An unidentified small, black, ant-like tenebrionid, Herpiscius sp. (Fig. 8K),
with glabrous elytra was very common in debris on the small foredunes skirting
the beaches along the west coast. The larvae feed on the roots of the dune plants,
and the white pupae (Fig. 8L) were usually found in the loose sand during the
summer and early autumn. One often sees these beetles running on beds in
houses and rondavels of holiday-makers near the beach, where they are generally
mistaken for ants or small spiders. Further away from the coast, they have been
found to visit dry cow-pats during the warm months, probably for the shelter
provided by the hard crust of the dung.
In Zimbabwe the matt, dark-brown to blackish Herpiscius sommeri Solier,
which, according to collection data, is widely distributed in the Cape Province,
was found to occur in large numbers among the stilt roots of the young maize
plants. It is rather similar to the previously mentioned unidentified species of
Herpiscius, but is easily distinguished by the distinct longitudinal striae on the
elytra. Herpiscius sommeri beetles were responsible for the destruction of the
anchor roots, causing severe damage to the maize plants (Rose 1964). Other
tenebrionids common in the debris under the loose sand on the dunes were
species of Phaleriderma and Gonopus, but their habits are unknown.
Dry cow-pats, particularly those that are fairly large, provide shelter for a
variety of tenebrionids, especially in open veld. The most common species found
during the surveys were the dusty surface-beetles, Gonocephalum arenarium
(Fabricius) and G. simplex (Fabricius), both of which occurred during the late
summer and winter months in most of the areas visited, often in association with
Blenosia exarata (Quensel), a dull-blackish beetle endemic to the Cape.
At least four species of mouldy beetles (subfamily Eurychorinae) were
collected under dry pats, of which the widely distributed Eurychora ciliata
(Fabricius) is the largest, measuring 13-14 mm in length. Lycanthropa plana
Haag, 6-7 mm in length, is more brownish and is endemic. Specimens of L. plana
which are indistinguishable from L. litoralis Koch were collected near Saldanha,
and may indicate that the existence of L. litoralis as a species as suggested by Koch
(pers. comm.) may not be valid. These beetles were all very plentiful along the west
coast. The genus Geophanus, another member of this subfamily, was represented
by a rather small, unidentified, black species (4-5 mm long). Although rather
scarce, it occurred in the same localities as the other members of this group.
Trachynotus reticulatus (de Geer) is blackish, elongate, 13-14 mm long, and is
characterized by a strong carina on each elytron. It was observed in association with
two other black forms, viz. T. acuminatus Quensel and an unidentified molurine
species; both of the latter species lack the rugosity of T. reticulatus. Several
Psammodes species were also found in greater abundance, but the common ones
were rarely seen under cow-pats. However, one large unidentified toktokkie,
about 20 mm long, occurred in the Karoo during the summer and autumn months.
SOUTH AFRICAN ARTHROPODS 233
The genus Hologenosis, which is rather similar to the previously mentioned
genus Zophosis, was represented by a dull-black, unidentified species of about
5,8 mm long; it was found to be quite common in the Karoo. A dorsoventrally
flattened, dark-brown species of Blastarnus (near pruinosis Fairmaire), with
seven to eight longitudinal carinae on the elytra, occurred together with the
species of Hologenosis. The related Emyon sp. (near brunnipes (Erichson)) found
with them, resembles a snout-beetle and is dark, castaneous brown, and of about
the same length as the Hologenosis sp.
The black, long-legged Stenocara dentata (Fabricius) (about 13 mm long)
occurred in the vicinity of Calitzdorp under dry pats devoid of any other insects.
Its body is covered with small tubercles and the shiny longitudinal costae on the
elytra are also tuberculate. The related S. Jongipes (Olivier) of about the same
size, its body with larger tubercles and lacking true costae, is another very
common species in the western Cape Province. It occurs under dry and even
semi-fresh cow-pats, often in association with the smaller and shiny, dark-brown
Ograbies subdentatus Koch and the dull dark-grey Asida unigena Peringuey, both
of which are endemic to the Cape. From observations it seems that under certain
circumstances these beetles feed on the damp particles of the dung. This also
seems to be true in the case of the shiny black Eutochia pulla (Erichson), which is
very similar to Alphitobius diaperinus (Fig. 11D), and which is widely distributed
in the Subsaharan region. Eutochia pulla may, however, be easily distinguished
from A. diaperinus by its somewhat longer, almost moniliform antennae,
better-developed striae on the elytra, and by the posterior margin of the pro-
notum, which is less sinuate on each side.
At least two greyish species of Cryptochile, both marked with brown patches
and also characterized by two strong carinae on each elytron, were found under
semi-dry to fresh cow-pats in the western parts of the Cape Province. They were
accompanied by a small, black species of Horatoma, whose presence was prob-
ably coincidental, and occasionally by a dull-black meracanthine, Acanthomera
dentipes (Fabricius).
The masses of stranded kelp on the intertidal and supratidal zones attracted
very few tenebrionids apart from the already mentioned Gonocephalum arena-
rium and Zophosis acuta, and the pale psammochromic Pachyphaleria capensis
(Castelnau), which feeds in both the adult and larval form on the stipe and fronds
of the large Ecklonia maxima, and probably also on Laminaria pallida and
Macrocystis angustifolia. According to Griffiths and Stenton-Dozey (pers.
comm.) this last-mentioned beetle forms one of the most important elements of
individual kelp strings.
Gonocephalum arenarium (Fabricius)
DESCRIPTION
Adult (Fig. 8A)
Dull, dark-brown to blackish species, covered with yellowish-brown to
greyish-brown adpressed hairs giving it a dark-greyish appearance. The two
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 8. Tenebrionidae, Anthicidae. A-B. Gonocephalum arenarium adult. A. Dorsal view.
B. Body setae enlarged. C. Gonocephalum simplex adult, body setae enlarged.
D-J. Gonocephalum arenarium \arva. D. Apical abdominal segment, dorsal view. E. Larva,
dorsal view. F. Larva, left lateral view. G. Head and first leg, left lateral view. H. Right
mandible, dorsal view. I. Left mandible, dorsal view. J. Front part of head, dorsal view.
K-L. Herpiscius sp. K. Adult. L. Pupa, ventral view. M. Anthicus apterus, adult.
SOUTH AFRICAN ARTHROPODS D359
species Gonocephalum arenarium and G. simplex (both found under cow-pats
in the western Cape) can be separated by the structure of the body hairs
(Fig. 8B—C), those of G. simplex being flattened, almost scale-like.
According to collection data in the South African Museum G. arenarium is
widely distributed in the Cape, South West Africa, east Africa, and is also present
on some of the islands along the Cape coast.
Larva (Fig. 8E-F)
Full-grown larvae are about 16 mm long and pale, yellowish red in colour;
young specimens usually with pale, yellowish-white, longitudinal line over dor-
sum of body. Claws simple, acute and rounded on dorsal side, but almost flat
ventrally and bordered on each side by a trenchant ridge, which extends nearly
from apex to base; spine at this point present on each side, posterior one rounded
and broad, other one longer and pointed. Front legs much more strongly de-
veloped than others. Mandibles (Fig. 8H—I) broad, only slightly longer than wide,
and clearly divided into cutting and molar areas; former longer in left mandible;
each cutting edge with preapical ventral tooth. Both molar areas pointed distally
when seen from above. Each mandible with lateral carina running from extreme
apex almost to middle, with long seta at this point on lateral face; also with dorsal
carina which is fairly rounded longitudinally and bearing two spines dorsally;
anterior spine short and rounded, posterior one longer and pointed. There are
also two short spines placed medially on labrum of G. arenarium (Fig. 8J), which
Jack (1918) does not record in G. simplex.
Jack (1918) describes the larva of G. simplex and it appears to be very similar
to G. arenarium, possessing almost the same number of spines on the ventral side
of the front legs (Fig. 8G) and thirteen to twenty-one spines on the anal segment
(Fig. 8D). They differ, however, in that G. arenarium has one small basal spine or
strong seta (or two in some cases) laterally on each pseudopod, which, according
to Jack, is absent in G. simplex.
BIOLOGY
The dusty surface-beetles are polyphagous and may attack both roots and
stems of various wild plants and vegetables. The larvae of G. arenarium have
been collected during October to December on the foredune system feeding on
the roots of the dune plants, the adult beetles emerging during January. The
beetles were found to feed on the leaves and stems of Arctotheca populifolia along
the west coast, causing feeding marks similar to those made by boring caterpillars.
Adults of this species were also observed under stranded kelp along the west
coast, just above the high-water mark.
Gonocephalum arenarium, together with the dull-black scarab, Trox horri-
dus Fabricius, was collected by members of the Percy FitzPatrick Institute of
African Ornithology of the University of Cape Town during their surveys on
Malgas and Marcus islands near Saldanha as well as on Dassen Island. Although
they describe the vegetation on Marcus Island as being rather sparse and
236 ANNALS OF THE SOUTH AFRICAN MUSEUM
G. arenarium is usually associated with plants, the material brought to the islands
by cormorants for nesting purposes may fulfil this beetle’s needs.
Pachyphaleria capensis (Castelnau)
DESCRIPTION
Adult (Fig. 9A)
Oval, pale straw-coloured to pale yellow. Some specimens with darker patch
in middle of elytra; head somewhat darker, in some specimens piceous. Finely
punctured, sparsely on elytra; latter with about seven weakly developed striae
each, smooth and fairly shiny. Pronotum on each side just in front of hind margin
with small, somewhat oblique impression. Length 6,6—7,5 mm.
Widely distributed along the Cape coasts and collected as far north as South
West Africa.
Larva (Fig. 9B—-C)
Subcylindrical or somewhat flattened; 13-16 mm long when fully grown,
pale straw yellow with darker or brownish patches on dorsum. Latter divided by
distinct ecdysial suture, which is fairly clearly visible over first four to six abdomi-
nal segments. Pronotum somewhat larger than other segments. Sparsely covered
with fine golden-brown hairs. Ninth abdominal segment (Fig. 9D-E) somewhat
wider than long, rounded in dorsal view, excavated in middle and bearing six to
eight strong, short spines on posterior border, which is slightly emarginate in
some specimens. Four long setae present dorsally near anterior margin. Pygopo-
dia small. Abdominal spiracles circular, those on first segment slightly larger than
others, usually visible on ventral side. Mesothoracic spiracle oval, larger than
those on abdomen. Apart from medium-long and shorter spine-like setae, integu-
ment is covered with minute spines, only visible under high magnification
(Fig. 9G). Legs (Fig. 9F) well developed, with reduced number of spines; those
on prothorax somewhat stronger; tarsungulus of each leg simple and acute and
with single ventral and posterolateral spine near base.
Head (Fig. 9H, J-K)
Cranium nearly twice as wide as long, with sides fairly convex and somewhat
emarginate. Brownish yellow to straw yellow in colour; frontal and coronal
sutures fairly distinct, frontoclypeal suture well demarcated. Very minutely re-
ticulate and fairly shiny, sparsely punctate, the punctures shallow; chaetotaxy as
illustrated. Hind margin ventrally deeply emarginate in middle; postgenae sepa-
rated by poorly demarcated gula. Neck absent. Antennae (Fig. 91) tubular,
second segment somewhat longer than first and wider distally, bearing some
minute setae apically; third segment small, much shorter than second and bearing
long seta apically as well as two or three smaller ones. Five darkly pigmented,
almost black ocelli present laterally, appearing as one eye when seen from
above.
SOUTH AFRICAN ARTHROPODS 257
Fig. 9. Tenebrionidae. Pachyphaleria capensis. A. Adult. B-U. Larva. B. Dorsal view.
C. Left lateral view. D. Ninth abdominal segment, dorsal view. E. Ninth abdominal segment,
left lateral view. F. Right mesothoracic leg. G. Integument, highly magnified. H. Head, dorsal
view. I. Antenna. J. Head, ventral view. K. Head, left lateral view. L. Epipharynx. M. Right
mandible, cephalic view. N. Right mandible, dorsal view. O. Labium, ventral view.
P. Hypopharynx. Q. Left maxilla, dorsal view. R. Hypopharyngeal sclerite enlarged. S. Left
maxilla, ventral view. T. Mala of left maxilla, cephalic view. U. Left mandible, dorsal view.
V-X. Pupa. V. Left lateral view. W. Comb-like structure on first and second abdominal
segments. X. Apex, ventral view.
238 ANNALS OF THE SOUTH AFRICAN MUSEUM
Labrum and clypeus
Labrum free, with anterior margin very rounded or convex and bearing six
strong spine-like setae dorsally, as well as three smaller setae on each side on
anterior margin. Clypeus almost trapezoidal and with four setae on stronger
sclerotized postclypeus.
Epipharynx (Fig. 9L). Tormae symmetrical; posterolateral processes short
and triangular median ones long and slender; the anterolateral extensions present
seem to be better developed and longer on right side, furnished with fine hairs
which form oval patch on each side of raised oblong central disc; the hairs on each
side extending on to central posterior area behind which are eight small spines or
sensilla; fine setae also extending on to anterior part (zz), forming a fairly large
brush (z) on anterior margin, also visible dorsally. Central disc with four large
anterior and four smaller posterior sensilla, as well as two short spines posterior
to these. Laterad of each posterior tormal process two spines present in most
- specimens examined. lateral margin anteriorly with three strong sickle-shaped or
lanceolate spines of which front one is apically rounded and somewhat more
strongly developed.
Mandibles (Fig. 9M-N, UV)
Similar to those of Gonocephalum arenarium (Fig. 8H-I). Broad, left
mandible slightly longer than wide, right one about as wide as long. Cutting
edge bidentate, lower tooth (tz) more rounded in some specimens; upper one
(t:) acute when seen from the dorsal side. Upper tooth (ti) on each mandible
extended dorsolaterally as trenchant ridge almost to molar area; latter with
broad almost rounded tooth (ts) dorsally and also extended ventrally as
broad tooth (ts); whole forming an almost triangular ridge on both mandibles
(Fig. 9M). On right mandible an extra rounded tooth (ts) present dorsally
about half-way between apex and molar area; on left one only an indication of
this tooth present. Lateral margin has a break at about middle, lateral face
sloping down from this break, with weaker sclerotization and bearing two strong
dorsal setae. |
Maxillae (Fig. 90, S-T)
Mala fairly broad, tapering to apex, with some fine, long hairs dorsally in two
longitudinal rows next to spines and some near lateral margin. Mesal margin with
two rows of spines or spine-like setae, curved at their tips, about twelve in dorsal
and nine in ventral row. Stipes ventrally with three long setae and two shorter
setae on lateral side, of which distal one is largest. Maxillary palp three-
segmented, with first two segments of about equal length, apical one shortest
and with about twenty sensory pegs at apex and minute seta in about middle on
mesal side; second segment with one ventral and one lateral seta, and some
minute spines near apex; first segment also with minute spines near apex and
short seta on external lateral margin near base. Cardo without setae.
SOUTH AFRICAN ARTHROPODS 239
Labium (Fig. 90)
Submentum somewhat trapezoidal, with two long setae. Mentum long, al-
most barrel-shaped or even tubular in some, with four setae. Prementum with two
setae. Ligula short, about half the length of first palpal segment and bearing two
apical setae and some minute spines on dorsal side. Palp two-segmented, with
segments of about equal length; apical one with about twenty sensory pegs at
apex; first segment with minute dorsal spines and a tiny lateral seta at base in most
specimens examined.
Hypopharynx (Fig. 9P, R) medially raised and convex from side to side and
conical, hypopharyngeal sclerite as short, broad, truncate tooth, almost triangular
in some specimens.
Pupa (Fig. 9V-X)
Whitish at first, but becoming yellowish later; about 6 mm long. Row of fine
golden setae present on front margin of pronotum, extending along anterior part
of lateral margin. About four setae present on head. ‘Gin-traps’ absent from
median area of abdominal dorsum, but each segment with lateral comb-like
extension (Fig. 9W) bearing four to six setae. Number of setae and fleshy
protuberances diminishing posteriorly. First seven pairs of spiracles clearly visible
laterally in fold formed by comb-like extensions; first two pairs of spiracles close
together and more or less covered by wings; eighth pair inconspicuous. Extreme
apex (Fig. 9X) has appearance of flat lobe, widening posteriorly, with two spines
on each corner, lateral ones shortest. Developing genital capsule takes shape of
two swellings; in some with extra projection of about same length laterad of each
conical swelling.
BIOLOGY
Pachyphaleria capensis beetles are found on the beach all the year round and
feed on decaying and fresh kelp. In areas where kelp is scarce they become
scavengers and feed on animal or bird carcasses on the beach. They were found in
wet sand under kelp at a depth of 150 to almost 300 mm. Larvae of this species were
collected along the west coast of the Cape Province during the autumn and early
winter and feed mostly on the fresh parts of washed-up kelp. When confined to a
small space or when food becomes scarce they become cannibalistic. As in the case
of the beetles, the larvae will feed on bird carcasses when kelp is not available.
There is a short prepupal stage of 2—4 days. Pupae are found in loose or hollowed-
out chambers in the damp sand and are usually hooked to the remains of the larval
skin. In the laboratory, pupal stages lasted about 16 days during May.
Tribolium destructor Uyttenboogaart
DESCRIPTION
Adult (Fig. 10A)
Dark-brown to piceous-brown, elongate beetle, 3,9—-5,3 mm long. Head and
pronotum fairly densely punctured; punctures large and oval; elytra each with
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 10. Tenebrionidae. Tribolium destructor. A. Adult. B-P. Larva. B. Dorsal view. C. Left
lateral view. D. Antenna. E. Right mesothoracic leg. F. Head, dorsal view. G. Head, ventral
view. H. Head, left lateral view. I. Labrum, dorsal view. J. Epipharynx. K. Left maxilla,
ventral view. L. Left maxilla, dorsal view. M. Hypopharynx. N. Labium, left lateral view.
O. Left mandible, dorsal view. P. Right mandible, dorsal view. Q-S. Pupa. Q. Ventral
view. R. Abdominal apex, ventral view. S. Comb-like structure on first two abdominal
segments.
SOUTH AFRICAN ARTHROPODS 241
nine to ten longitudinal striae and with row of large shallow punctures between
striae, lateral ones more conspicuous; area between punctures very finely reticu-
late and whole insect therefore fairly shiny. Only legs and antennae covered with
fine yellowish-brown hairs.
Tribolium destructor belongs to the confusum-group (Hinton, 1948), but may
easily be distinguished by its larger size and the fact that the narrowest part of the
eye, where it is divided by a carina on the side of the head, is usually as broad as
two facets of the eye, and the frons is beset with punctures that are as long as the
eye facets. In T. confusum, the narrowest part of the eye is no broader than one
facet.
This species is widely distributed in the Republic of South Africa according
to collection data and often appears in stores and butcheries in the western Cape.
It is a native of Africa and is also a pest in North America (Hinton 1948).
Larva (Fig. 10B—C)
General body colour creamy white or whitish yellow, most segments brown
dorsally; the head, thoracic and last two abdominal segments darker. Ecdysial
Suture distinct. Length 9,5-10 mm when fully grown. Body subcylindrical to
almost cylindrical and sparsely covered with long golden setae, more or less
arranged in two rows on each segment. Urogomphi short, acute, divergent and
directed upwards. Spiracles circular, visible laterally; mesothoracic pair very
slightly larger than others. The pygopodia can be retracted into the ventral
aperture of the ninth segment.
Legs (Fig. 10E) well developed, all of about equal length and with a reduced
number of setae. Tarsungulus acute, simple, without spines and slightly shorter
than length of tibia. Dorsal integument very similar to that of Pachyphaleria
capensis, with long setae and short spines, but lacking minute spicules.
Head (Fig. 10F—H)
Cranium only slightly wider than long, minutely reticulate, only slightly shiny
and also somewhat rugulose, particularly the frons. Frontoclypeal suture well
developed: frontal and epicranial sutures inadequately indicated. Antennae
(Fig. 10D) three-segmented, second segment about twice as long as first, with
small sensory tubercle on anteroventral side as well as at least one or two minute
spines below it and one situated dorsolaterally; third segment small, about as long
as first, with a medium long apical seta. One or two pigmented eye-spots visible
behind antennal base, but separate ocelli not demarcated in specimens examined.
Postgenae ventrally separated by fairly wide gula. Neck absent. Setation as
illustrated; some setae very long. Head capsule, particularly side and ventral part.
with numerous small spines.
Labrum and clypeus
Labrum (Fig. 101), which is partly concealed by clypeus when viewed from
above, is characteristic of this species. Clypeus about three times wider than long,
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
its anterior margin almost straight or slightly concave in most specimens seen and
bearing two medium-long setae on each side. Labrum much shorter than clypeus
and slightly more than twice wider than long; setal pattern as illustrated.
Epipharynx (Fig. 10J). Under high magnification four large sensory spots or
cones visible on raised median area and just posterior to these a transverse row of
four minute sensory cones present, as well as two small spines. Most specimens
examined also with two large sensory cones near anterior margin, just posterior to
paramedian setae. Setation of each side of median area consists of minute
spicules, which are either simple or with two to four teeth. Tormae symmetrical,
on each side with oblique row of fine hairs as well as two sensory cones and group
of eight tiny sensory spots or spines between posterior arms. Anterior margin
with four short setae and three large almost sickle-shaped setae posterior to these
on each side.
Mandibles (Fig. 10O-P)
Somewhat longer than wide, with break on lateral margin not so obvious as
in Pachyphaleria capensis. Each mandible with two setae situated on short,
oblique rounded dorsal carina, as well as one long anterior and two posterior
setae on lateral face; latter setae situated ventrally near base. The median tooth
(t3) on right mandible much less pronounced than in P. capensis.
Maxillae (Fig. 1OK-L)
Mala devoid of setae on ventral side, except one spine-like seta near base in
some specimens; dorsally, however, with some long hairs and mesal margin with
two rows of spines, seven to ten on each side (as in P. capensis). Stipes with three
long ventral setae, one seta on lateral side near palpifer and about two short setae
near cardo; dorsally devoid of setae. Cardo without setae, except for one or two
very short ones in some specimens. Maxillary palp three-segmented; segments
short; first two nearly of equal length; the second with one lateral and one ventral
seta; apical segment somewhat longer and with some sensory cones at apex and
minute seta on mesal margin some distance from apex.
Labium and hypopharynx (Fig. 1OM-N)
Similar to that of P. capensis (including ventral setae), but ligula absent and
mentum shorter. Hypopharyngeal sclerite is more oval, appearing cup-shaped
and in most specimens examined with roughly M-shaped sclerotized band or base
posterior to it. Glossa with four setae between and posterior to two-segmented
palps. Surface of glossa and area around hypopharyngeal sclerite covered with
minute spines. In lateral view submentum is raised above level of mentum. Labial
palp with apical segment somewhat longer than first and with seta on external
margin near base.
Pupa (Figs 10Q-S, 12B)
Length about 5 mm; at first pale or whitish except for apical spines, which
are brownish or yellowish. It gradually darkens in colour and just before imago
SOUTH AFRICAN ARTHROPODS 243
emerges it becomes piceous. The pupa of Tribolium destructor is rather similar to
that of P. capensis (see Fig. 9V—X), but is more elongate, with somewhat more
hairs on the body, particularly on abdomen. It differs from the latter by smaller
size and longer apical spines. Developing genital capsule (Fig. 10R) takes shape
of two fairly long diverging, conical projections, which in the case of P. capensis
are placed more parallel. Lateral extensions of abdominal segments (Fig. 10S)
each with two conical tubercles and two to three setae (at least three to four in
P. capensis); anterior and posterior teeth strongly developed. Spiracles similar to
those of P. capensis.
BIOLOGY
Larvae of Tribolium destructor were collected during September in butch-
eries around Paarl and Wellington in sawdust sweepings containing small bits of
meat and dust. Their life-span was fairly long, at least 4-6 months, and pupae
formed during October to December produced beetles after 12—23 days. During
the winter months the pupal stage was somewhat longer. In most cases pupation
occurred in the sawdust mixture without the construction of pupal cells.
The newly-emerged beetles have yellowish-white elytra, with pale yellowish-
brown head, thorax and legs. After a day or two they become castaneous brown
and may remain pale coloured for up to 8 days, after which they attain their
normal piceous or brownish-black colour.
Alphitobius diaperinus (Panzer)
DESCRIPTION
Adult (Fig. 11D)
Blackish-brown to almost black, shiny, slightly oval beetle, whole dorsal
surface of head and pronotum finely punctured; each elytron with about six
weakly developed, longitudinal, punctured striae with some punctures in between
striae. Hind margin of pronotum sinuate on each side, antennae short, about as
long as length of pronotum, incrassate towards apex. Legs short. Length about
6 mm.
Cosmopolitan (Andres 1931; Cotton 1941) and commonly found in South
Africa.
Larva (Fig. 11A)
Subcylindrical and very similar to that of Pachyphaleria capensis (see
Fig. 9B), creamy white, with brownish coloration on tergal plates, ecdysial suture
as thin whitish line along middle of dorsum, only faintly visible on second to last
abdominal segments in most specimens seen. Pronotum somewhat larger than
other segments as in latter species; whole body sparsely covered with fine golden
hairs. Ninth abdominal segment (Fig. 11B) convex, triangular when seen from
above; apex ending in an upcurled spine; laterally and dorsally with about sixteen
spine-like setae, also with some long slender setae, particularly on ventral side;
dorsally also four long setae in transverse row in about middle of segment.
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 11. Tenebrionidae. Alphitobius diaperinus. A-C. Larva. A. Left lateral view. B. Ninth
abdominal segment, dorsal view. C. Right mesothoracic leg. D. Adult. E—O. Larva. E. Head,
dorsal view. F. Head, ventral view. G. Head, left lateral view. H. Epipharynx. I. Left
mandible, dorsal view. J. Right mandible, dorsal view. K. Left maxilla, dorsal view.
L. Labium, ventral view. M. Hypopharynx. N. Left maxilla, ventral view. O. Hypopharyngeal
sclerite, highly magnified.
SOUTH AFRICAN ARTHROPODS 245
Pygopodia fairly well developed and usually seen as two conical processes
below ninth segment. Spiracles circular, that on mesothorax the largest; first
abdominal spiracle also somewhat larger than other abdominal spiracles. Legs
(Fig. 11C) well developed, of about equal size and with reduced number of spines
and setae; tarsungulus simple, fairly long, acute and with single ventral seta.
Dorsal integument as in P. capensis, including the short spines and spicules.
Length about 11 mm when fully grown.
Head (Fig. 11E-G)
Cranium nearly twice as wide as long, sides convex; shiny, somewhat sha-
greened, golden yellow; central transverse area darker, almost reddish brown in
some specimens, including clypeus and labrum. Mandibles lighter in colour than
rest of head. Coronal and frontal sutures clearly indicated, latter branched near
antennal bases (apparently also the case in Pachyphaleria capensis and Tribolium
destructor, but not as distinct as in this species). An elongate blackish eye-spot
present behind antennal bases, but individual ocelli not demarcated in all speci-
mens examined. Chaetotaxy as illustrated. Hind margin dorsally and ventrally
only slightly emarginate in middle, postgenae separated by fairly well-indicated
gula. Antennae as in P. capensis, second segment about one-third longer than
first; third segment also small, less than one-third the length of second and
bearing long apical seta.
Labrum and clypeus
Labrum about twice as wide as long and somewhat narrower than clypeus, its
front margin almost straight, lateral margins convex. About ten slender setae
present of which two are situated on central area and two in middle near anterior
margin; also four short setae present along front margin. Clypeus asin P. capensis,
slightly less than twice as long as labrum, with four setae on postclypeus.
Epipharynx (Fig. 11H). Rather similar to that of T. destructor (see Fig. 10J).
Tormae symmetrical, posterolateral processes short and broad, median ones long
and slender. Fine spine-like setae cover most of elongate area on each side of
raised central part and forming an almost single row along front margin, the setae
longer on tormae and along each side of central part, which bears six large sensilla
and transverse posterior row of four smaller ones; behind these two spines
present; also two longer spines or teeth present between posterior arms of tormae
near transverse central bar, and just behind right tooth a row of about six longer
spinules (or setae). Eight spines (or sensory spots) present on central area just
behind long posterior teeth as well as one on left side between posterolateral
process and median arms, and two in same position on right side. Three sickle-
shaped setae present on each lateral margin of which the distal one is more
strongly developed than others (rather similar to those of P. capensis).
Mandibles (Fig. 111-J)
Similar to those of P. capensis (see Fig. 9M-N, UV), fairly broad with
flattened, almost triangular lateral face, bearing about six slender setae on its
246 ANNALS OF THE SOUTH AFRICAN MUSEUM
upper and lower margins as indicated in drawing. Cutting edge bidentate on both
mandibles with lower tooth (tz) smaller than upper or apical one. Molar area with
upper tooth (ts) rounded on left mandible, but more acute on right one. Lower
molar tooth (ts) fairly acute on both mandibles. A small, distinct, rather acute
tooth (ts) present on trenchant ridge from upper scissorial to upper molar tooth,
close to latter on left, but in about middle on right mandible (in both P. capensis
and 7. destructor this tooth is almost obsolete in left mandible).
Maxillae (Fig. 11K, N)
Similar to those of P. capensis, broad, mala with double longitudinal row of
about nine spines on mesal side, dorsally beset with fine slender setae; ventrally
devoid of slender setae. Stipes ventrally with three long setae of which one is
situated near base, just anterior to cardo, which is fairly well indicated and
without setae. Also small spine or spine-like seta present near mesal margin;
dorsally stipes without setae, except one long and two short setae placed some-
what dorsolaterally. Maxillary palpi three-segmented, first two segments of about
equal length, apical one somewhat shorter and with short subapical seta on mesal
side and about twenty sensory pegs on apex; first segment with short dorsolateral
seta near base and second segment with two very short ventral setae and one long
lateral seta near apical border. Palpifer without setae and forming small, almost
ring-like segment at apex (not observed in the specimens of the other two
previously mentioned species).
Labium (Fig. 11L—M)
Submentum broad, oval, with two setae, one on each side near base of oval
convex sclerite of cardo. Mentum longer than wide, somewhat barrel-shaped, its
sides convex and bearing four setae on sclerotized posterior part. Prementum
shorter than mentum with two long median setae near apex and short seta placed
ventrolaterally; also minute spines posterolaterad of each median long seta.
Glossa without setae as in P. capensis, ligula short, about three-quarters the
length of first palpal segment and with two setae at apex. Labial.palp two-
segmented, first segment broader than long and somewhat longer than apical
conical segment and with two short ventral setae near apical border one mesally
and one laterally; second segment with about twenty sensory pegs at apex.
Hypopharynx (Fig. 11M, O). Medially raised, the hypopharyngeal sclerite
appearing cup-shaped; however, when viewed slightly from the side its dorsal
aspect is an almost inverted horseshoe-shaped sclerite with a somewhat dome-
shaped ventral side.
Pupa (Fig. 12A, C-D)
Rather similar to that of Tribolium destructor (see Fig. 10Q—R) but larger,
measuring 6,6 mm long and 2,7 mm broad and lacking the long setae on pro-
notum; latter broad, about twice as wide as long (in 7. destructor only slightly
wider than long). Head, pronotum, wing-covers and pleural surfaces with some
SOUTH AFRICAN ARTHROPODS 247
mm
Fig. 12. Tenebrionidae. A. Pupa of Alphitobius diaperinus, left lateral view. B. Head and
pronotum of pupa of Tribolium destructor, left lateral view. C-—D. Pupa of Alphitobius
diaperinus. C. Abdominal apex, ventral view. D. Comb-like structures on first two abdominal
segments, dorsal view.
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
fine setae, also some setae present on legs and mandibles. Thorax and abdominal
terga covered with minute tubercles, those on thorax larger than the others (in
T. destructor the body surface is more shagreened or wrinkled). Spiracles similar
to those of 7. destructor and P. capensis, six pairs clearly visible, seventh pair
indistinct. Developing genital capsule and lateral extensions of abdominal terga
including anterior and posterior teeth similar to those of 7. destructor; however,
the dentate areas more elongate.
BIOLOGY
The genus Alphitobius belongs to the subfamily Ulominae, the members of
which are mostly detritivorous. Only about ten species of this genus occur in the
Subsaharan region, one, A. /eleupi Koch, being commonly found in the nests of
gerbilles (Tatera species—Koch 1953). As far as is known only four members
have been recorded from southern Africa, viz. A. hobohmi Koch from South
~ West Africa, A. karrooensis Koch from the Cape Province, A. ulomoides sensu
Koch (nec ulomoides Solier, which is a native of Chile), and the widespread
A. diaperinus, which has become a pest of stored grain and flour. However,
according to Cotton (1941) it does not damage sound and dry grain. There is also
a possibility that this latter species plays a role in the transmission of acute
leukosis in chickens (Eidson et al. 1965).
The lesser meal-worm A. diaperinus often occurs, together with skin-and-
hide beetles, during the third or dermatophagous stage of decay of carcasses and
cadavers and is commonly found in fowl manure, particularly in the closed-type
poultry batteries with a fairly high and constant temperature of about 20 °C.
Where very large populations of this beetle and of Dermestes maculatus occur the
beetles compete with the normal fly species to such an extent that such poultry
houses are almost free of the common house-flies, false stable-flies and lesser
house-flies. Under these circumstances larvae of the skin-beetle may become
troublesome in their effort to secure suitable space for pupation and may cause
severe damage to roof timbers, door frames, etc. Application of an insecticide
such as carbaryl in a broad band around the inside walls and the floor adjoining
the walls of the building has controlled the wandering larvae of this pest in the
past, whilst allowing the meal-worm larvae to continue feeding in the manure;
however, large populations of the latter may have a depressing effect on hens
kept on floor litter. When crowding occurs in the case of large populations larvae
of both species become predacious and even cannibalistic. Young larvae are
preyed upon by the histerid Carcinops minuta (Fahraeus), breeding in the
manure.
Fairly young larvae of the lesser meal-worm collected on a decaying turtle
carcass during February lived for at least 30-40 days in the laboratory and
moulted several times before pupation occurred; the pupal stages lasted for about
10 days (temperature about 25 °C). According to Barké & Davis (1969) the pupal
stadium was 7-11 days at about 21 °C and 70 per cent relative humidity, and the
total life-cycle 69-91 days; the eggs hatched over a period of 6—10 days. Cornwell
SOUTH AFRICAN ARTHROPODS 249
(1973) gives the minimum development period as 40 days at 25 °C. When freshly
formed, the pupae are creamy white but later change to a darker colour. The
newly-emerged beetles are pale brownish.white and assume their normal colour
in a day or two.
SERIES SCARABAEIFORMIA
Family Scarabaeidae
Small to large, usually stout-bodied or robust beetles, sombre to brightly
coloured with partly deflexed head and antennae with lamellate club of three to
seven segments; elytra usually exposing pygidium; wings well developed, absent
in some forms such as certain dung-rollers and trogids (the latter often regarded
as a separate family, Trogidae). Head and pronotum often with horn-like pro-
cesses; front tibiae fossorial; tarsal formula 5—5—S.
This is a large family composed of two groups. In the first at least four sub-
families are involved and the members are either saprophagous or fungivorous.
This group includes the true dung-beetles or dung-rollers of the subfamily Scara-
baeinae (Coprinae) in which the adults have eight- to nine-segmented antennae
and the larvae have a distinct dorsal hump (Fig. 19C). Their legs are usually
reduced or the claws are absent. Both adults and larvae are found in dung and
carrion and the adults may even feed on fungi. This subfamily includes fairly large
and robust beetles, having a body length of up to 57 mm in Subsaharan forms,
such as Heliocopris gigas (Olivier), ranging from Zimbabwe to tropical Africa. It
also includes the flightless, ball-rolling Circellium bacchus (Fabricius), which has
been recorded from the Cape Province, Transvaal, and Mozambique (Ferreira
1967) but according to Tribe (1978) is now restricted to the eastern Cape Prov-
ince, particularly the Addo Elephant Park, and may be heading for extinction. It
is most probably a forest-bound species and may still be present in fair numbers in
such localities in Mozambique, Transvaal, and the north-eastern parts of southern
Africa.
Some of the smaller members of this subfamily such as the widely distrib-
uted, bronze-and-green dung-beetle, Onitis aygulus (Fabricius), and the black
O. caffer Boheman do not roll balls but burrow into the soil beneath animal
droppings and utilize the contents below the crust of the cow-pats for oviposition.
These are strong fliers and powerful diggers and prefer fresh dung, particularly
during the winter months.
Three species of the smaller genus Euoniticellus were collected from fresh
dung during the surveys: E. intermedius (Reiche), E. triangulatus (Harold) and
E. africanus (Harold), of which the first two occurred in large numbers almost
throughout the year in most of the areas visited. The indigenous E. africanus
(Fig. 14F) was only occasionally encountered, usually in the Montagu area and
along the south coast during the summer and late spring. These beetles are dark
brown with black marks and spots on their bodies, and vary in length from
7-10 mm in E. intermedius and E. triangulatus to 10-11 mm in E. africanus.
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
Oniticellus pictus (Hausmann) (Fig. 18A) and O. planatus Castelnau
(Fig. 19A) also visit fresh dung, as their pupal cells were often found in large
numbers in semi-fresh pats in the Robinson Pass and near Mossel Bay during the
autumn. Oniticellus pictus resembles the members of the genus Euoniticellus, but
is easily separated from them by the clearly demarcated black marks on its body.
Both O. pictus and O. planatus were found in the same pats.
Chironitis scabrosus (Fabricius) was very common near Montagu and in the
Karoo during the summer. This beetle is medium-sized, castaneous or testaceous
brown with a bronzy tinge, having the pronotum with irregular rugae and the
elytra with blackish tubercles. In fresh cow-pats it was usually associated with
C. hoplosternus (Harold), which is rather similar, but has the pronotum some-
what wider, is more greenish and is without the rugosity.
The black species of Copris and Onthophagus were observed in small num-
bers on fresh dung in various localities during the spring and winter, Copris
~ anceus Olivier (Fig. 13A) being the most common. Males of C. anceus
(19-20 mm long) have a triangular horn on the head; females, which are some-
what smaller, have the pronotum slightly convex, without horns and the head
with only a short tooth-like horn.
Onthophagus minutus Hausmann (about 6 mm long), black in colour, with a
short tooth-like horn on the head of the male, was occasionally seen feeding on
fresh and semi-fresh dung near Mamre and other Sandveld areas. Another
species, O. immundus Boheman (matt black and 5—5,8 mm long), was frequently
found in Heidelberg during the spring and summer. The males are easily recog-
nized by two parallel horns on the head. In this area it was associated with the
much larger (9-12,6 mm) O. binodis Thunberg (Fig. 14A—B), also matt black.
Males of the latter have a rectangular, carinated, raised lobe in the middle of the
pronotum. This species also occurred commonly in the Karoo during the autumn.
Onthophagus cameloides D’Orbigny (Fig. 13E), dull black and about 10 mm
long, appeared in partly dry dung in the western and northern parts of the Cape
Province, but was not numerous.
A rather peculiar, dark-brown scarab, Drepanocerus kirbyi Kirby (Fig. 14D),
about 6mm long with three long horns, one on the head and two on the
pronotum, seems to be fairly common in the Robinson Pass where it was found
feeding on fresh dung on several occasions during the summer and autumn
months. The large, shiny black dung-roller, Neateuchus proboscideus (Guérin)
(Fig. 14G) (almost 30 mm long), on the other hand, was collected only occasion-
ally during the spring in the north-western parts. Other small scarabs belonging to
this subfamily, found in fresh to semi-fresh cow-pats, were the greenish-black
Epirinus aeneus (Wiedemann) and the larger, blackish E. flagellatus (Fabricius)
(Fig. 13D), both endemic to the Cape according to collection data.
The small dung-beetles of the subfamily Aphodiinae include the regular
inhabitants of herbivore droppings that never bury any dung but feed on it as long
as enough food is available. The larvae of these coprophagous beetles with their
SOUTH AFRICAN ARTHROPODS ide |
a
of
A F
x
Fig. 13. Scarabaeidae. A. Copris anceus. B. Trox fascicularis. C. Unidentified species of
Trogositidae. D. Epirinus flagellatus. E. Onthophagus cameloides.
US ANNALS OF THE SOUTH AFRICAN MUSEUM
nity,
~~
Fig. 14. Scarabaeidae. A. Onthophagus binodis, 6. B. Onthophagus binodis, head and pro-
notum of 2. C. Trox horridus. D. Drepanocerus kirbyi. E. Temnorrhynchus retusus.
F. Euoniticellus africanus. G. Neateuchus proboscideus. H. Eucamenta castanea. 1. Adoretus
ictericus.
—<—_ TO OETETETET_E_E_EEO ee eileen
SOUTH AFRICAN ARTHROPODS 253
well-developed mouth-parts are able to feed on even fairly dry dung particles,
whereas in the case of the adults, as in other smaller members such as the previously
mentioned genus Oniticellus, the mouth-parts are more membraneous and they are
therefore compelled to feed on moist or soft dung. This is also substantiated by
their intestinal contents which generally only contain soft liquid compounds.
In the adults the antennae are usually nine-segmented with three-segmented
club and the larvae have well-developed legs and lack the dorsal hump. A few
species are phytophagous (not present in South Africa) and at least two species,
Aphodius lividus (Olivier) and A. granarius (Linnaeus) (both present in South
Africa), are cosmopolitan.
Some 150 species of aphodiines have been recorded from southern Africa of
which about ninety-six belong to the genus Aphodius. These small scarabs are
rather important as they play a vital role in the disposal of animal dung in the
veld. During the surveys at least ten species were found to be attracted to fresh
droppings, and some used the dung as food. Aphodius procerus Harold, A. consi-
milis Boheman and A. Jaetus Wiedemann were found to be truly coprophilous,
actually breeding in this medium; A. consimilis and A. badius Boheman were
found even after the droppings had dried out to some extent. Aphodius badius,
which is rather similar to another small, dull, dark-brown scarab Drepanocanthus
lineatus (Wiedemann), but much smaller, has in addition been collected in fresh
dung on the beach along the west coast of the Cape Province together with the
previously mentioned Epirinus aeneus.
Most of the Aphodius species are widely distributed: A. procerus (Fig. 16A)
and A. maculicollis Reiche occurred more commonly to the east of the Cape
Peninsula and in the Karoo, while A. hepaticus Roth was found only in June in fresh
dung in the Little Karoo. All the other species occurred almost throughout the
year, A. consimilis being particularly abundant during the early spring. This species
is often attracted to lights in houses, especially during the late summer and autumn.
The larvae of A. /aetus and A. procerus were fairly heavily preyed upon
during the winter and early spring by the larvae of a black and white-banded
dexiid, Pretoriamyia sellifera van Emden, which occurred throughout the survey
area. The larvae of an unidentified muscid of the genus Helina were also respon-
sible for the destruction of various small scarab grubs, mostly Aphodius species,
and it seemed to have the same habits and distribution as the species of
Pretoriamyia.
Puparia of Helina spp. can easily be separated from those of Pretoriamyia by
the small pale posterior spiracles placed close together at the hind end. In
Pretoriamyia the spiracles are larger and black and situated on the dorsal side of
the hind end.
Another aphodiine found in large numbers in fresh dung is the above-
mentioned Drepanocanthus lineatus. It was collected in almost all areas during
winter and spring; in the northern parts it appeared together with Coptochirus
excisus Harold, which is much smaller and more shiny. Along the south coast it
was found in association with Harmogaster intrusa (Peringuey) a small, dull,
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
dark-brown species with black pronotum. Coptochirus pallidipennis Harold, and
C. brachypterus Harold, were found to be very numerous in the Mamre area
during September when large numbers of both were seen to copulate. Copto-
chirus emarginatus (Germar), a very common species in fresh and partly fresh
dung all along the west coast almost throughout the winter and early spring, often
occurs in large numbers in factories manufacturing materials that contain
feathers, such as quilts.
All the Coptochirus species mentioned here are small (4,6—5 mm long), and
are recognized by the emarginate anterior border of the head. Coptochirus
emarginatus, C. pallidipennis and C. excisus are light brown, with the pronotum
and head a darker brown in the middle. The first-named species: also has dark-
brown patches on the elytra. Coptochirus brachypterus on the other hand has the
pronotum and head almost black in the middle and the elytra in the specimens
examined are darker than in the other three species.
A large number of small scarabs were still present in partly dry cow-pats that
were examined and of these the following were found to be prevalent: Aphodius
moestus Fabricius, A. bidentulus (Harold), A. lugubris (Boheman), Oniticellus
pictus, Coptochirus pallidipennis, C. brachypterus, Onthophagus cameloides,
Harmogaster exarata Harold, the two scarabaeines, Odontoloma dentinum
(Harold) and O. pygidiale Peringuey, Aphodius discoidalis Boheman and some-
times also A. badius, A. rubricosus Boheman, and the black maize-beetle,
Heteronychus arator Burmeister (subfamily Dynastinae). The last two species
even occurred when the pats were very dry and their crust hard and firm, while
A. moestus also bred in fresh and semi-fresh pats.
All the Aphodius species are more or less dark brown, but A. moestus
(Fig. 1SE) has lighter elytra, marked with short, longitudinal, black stripes. It is
widely distributed in South Africa and its larvae are most often found in associa-
tion with those of A. consimilis. Aphodius lugubris on the other hand seems to be
more restricted to the west coast and appears in rather small numbers throughout
the spring and autumn; it is commonly found to breed in the decaying reeds in
thatched roofs of houses and rondavels. The shiny-brown Aphodius discoidalis,
almost of the same size as A. moestus (about 7 mm), was collected in fairly large
numbers near Melkbos during March, whereas A. bidentulus (4,5-5 mm long),
which is slightly shiny and black, was found in the Heidelberg area of the Cape
Province during the early winter in semi-fresh to almost dry cow-pats.
Oniticellus pictus, already mentioned in connection with fresh dung, was
collected in almost dry pats near Hermanus and further east along the south
coast. Oniticellus militaris Castelnau (about 9 mm long, dull dark brown, with
rather indistinct black markings on the elytra), was found to be common in the
autumn in semi-fresh and also fairly fresh cow-dung near Mossel Bay and in the
Robinson Pass. Coptochirus pallidipennis and the almost black Harmogaster
exarata were occasionally found in very large numbers in fairly fresh as well as in
partly dry dung in most of the areas investigated. The latter is easily distinguished
from C. pallidipennis by the structure of the striae on its elytra. Of the species
SOUTH AFRICAN ARTHROPODS 255
invading fresh dung, Aphodius consimilis and A. procerus together with Onitis
caffer remained in the dried-out dung for fairly long periods.
Odontoloma dentinum (3,4 mm long), dull black in colour with striae similar
to those of the previously mentioned Onthophagus cameloides, and Odontoloma
pygidiale, similar to O. dentinum, but lacking the longitudinal, smooth line on the
anterior part of the pronotum, were collected together with other small scarabs in
semi-fresh dung near Stilbaai during the autumn and near Elands Bay during
midwinter. Both species seem to be restricted to the southern part of southern
Africa according to collections that were made in the past.
Usually the larvae of only one species were found in a single cow-pat; in the
case of Aphodius laetus, A. moestus and A. consimilis, however, the larvae were
collected in the same pats.
The other two smaller subfamilies belonging to this group of scarabs, the
Geotrupinae (with three tribes of which only one, the Bolbocerini with about
forty species, is represented in southern Africa) and the Troginae (with one
Subsaharan genus comprising forty-two southern African species), are often
placed as separate families.
Only three trogids were collected during the surveys, the largest being the
previously mentioned Trox horridus Fabricius (Fig. 14C), 17-18 mm long with
penicillate tubercles as well as serrate lateral margins. It was often observed
during the winter months on or in the vicinity of dry cow-pats near Saldanha. It
differs from 7. fascicularis Wiedemann (Fig. 13B), which was occasionally
attracted to partly-fresh dung, by its size and by the absence of wings. This latter
species is also dull black and widely distributed in the Cape Province and Natal.
Another small species, T. rhyparoides Harold, occurred only in decaying car-
casses. The larvae of one of the largest members of this subfamily, 7. procerus
Harold, not present in southern Africa, but widely distributed in the arid parts of
north Africa and Arabia, have been observed by Van Emden (1948) to feed on
the eggs of the desert locust, Schistocera gregaria (Forskal) in Somalia.
The second group of scarabs includes the major phytophagous members,
better known as chafer-beetles, cockchafers, June beetles, Christmas beetles,
monkey-beetles, fruit-beetles and rhinoceros-beetles. Some of them often swarm
in large numbers during the spring, defoliating fruit and forest trees; a few
species, however, are predacious. The larvae are typically scarabaeiform as in the
first group, C-shaped (except in certain members such as the fruit-beetles),
whitish, with well-developed legs increasing in length from front to back, and with
pale reddish-brown head, either with or without ocelli. They are known as white
grubs and feed on dung, other decaying organic material, or on the roots of
plants, and may cause severe damage to cultivated crops and lawn grass.
The rhinoceros-beetles (subfamily Dynastinae) contain about sixty southern
African species, mostly dark brown to black in colour, with ten-segmented
antennae. The most common and widespread member is probably the shiny
256 ANNALS OF THE SOUTH AFRICAN MUSEUM
dark-brown Oryctes boas (Fabricius), 35-40 mm long, with a long curved horn
on the head of the males. One of the largest Subsaharan species is Dynastes
centaurus (Fabricius), measuring about 55 mm long and occurring also in the
north-eastern parts of South Africa. (Males of the large Hercules beetle
D. hercules (Linnaeus) of Central America and the West Indies measure up
to 130 mm, including the horn.)
In the larvae of this subfamily, which generally feed on roots or other
decaying vegetable matter, the labrum is usually asymmetrical and the ninth and
tenth abdominal terga are completely fused (Crowson 1967). This is apparently
not the case with all the members of this subfamily (see description of the larva of
Temnorrhynchus retusus (Fabricius), p. 290). In the case of the previously men-
tioned black maize-beetle, Heteronychus arator, commonly found under semi-dry
to dry cow-pats, the larvae do very little damage; the adults, however, may cause
severe damage to young maize plants, trees, etc., and even lawn grass. The larger
H. licas (Klug), widely spread in the Subsaharan region, including the north-
eastern parts of South Africa, often causes considerable damage to maize and
sugar-cane in Mozambique (Jerath 1966), and in this case both larvae and adults
are responsible. Heteronychus consimilis Kolbe, which is of about the same size as
the black maize-beetle, is a serious pest of wheat in Kenya (Le Pelley & Goddard
1952); according to collection records it does not occur in South Africa. The
larvae of a few species are found in termite mounds and feed on the material of
which the mounds are made.
A regular visitor on the open beach along the west coast is the piceous-red
Temnorrhynchus retusus (Fig. 14E), rarely seen because of its habit of burrowing
into the sand, most often just above the low-water line. Larvae of this species feed
on plant roots and other organic matter and are common in sandy soils through-
out the year in certain parts of the Cape Peninsula. It is also present on Dassen
Island. The larvae of a related species, T. coronatus (Fabricius), somewhat bigger
than 7. retusus (about 23 mm long) and widely spread in the Subsaharan region
including South Africa, sometimes damage the underground parts of strawberry
plants (Oberholzer 1963).
Some 270 species of fruit-beetles (subfamily Cetoniinae) have been recorded
in southern Africa, the largest members found in the Republic of South Africa
being the black and light-grey Goliathus albosignatus Boheman (45-55 mm long)
and the green and white Dicranorrhina derbyana Westwood (35-50 mm long).
This subfamily includes the large, robust, maroon-coloured Goliath beetle
Goliathus goliathus (Drury), of tropical Africa, which may reach a length of
90 mm or more.
The larvae of these beetles usually feed on humus, particularly in compost
heaps, dead logs, etc., and are not curled like the other scarabs, but are able to
stretch their bodies and walk horizontally or even slide in a worm-like motion on
their backs. The ninth and tenth abdominal terga are completely fused as in the
dynastids, but the labrum is always symmetrical (Hayes 1929).
SOUTH AFRICAN ARTHROPODS 257
The adults, with ten-segmented antennae, are diurnal and usually feed on
nectar. Members of the genera Pachnoda and Rhabdotis are particularly fond of
ripening fruit and may cause considerable damage in orchards. The genus Rhino-
coeta includes three species that generally fly at night and are coprophagous.
Rhinocoeta cornuta (Fabricius) is the most wide-spread, its larvae being common
in partly fresh cow-pats along the south coast. Another typical member is
Trichostetha fascicularis (Linnaeus), about 23 mm long with green elytra and
black prothorax having four thin white longitudinal lines. It is common in proteas,
particularly the king protea (Protea cynaroides), often in association with the
smaller brownish T. capensis (Linnaeus), the larvae of which are also known to
cause destruction to thatched roofs in the Cape Peninsula, and T. signata
(Fabricius) whose larvae are occasionally found in dassie dung in the same
area. The larvae, when full grown, construct a fairly hard earthen cocoon in which
the pupae are formed. Immature stages of T. capensis have been observed in
the mounds of termites (Termes spp.), sometimes together with those of the
large dark-brown dynastid, Pseudocyphonistes corniculatus (Burmeister) (adults
30—40 mm long). They both feed on the material of which the mounds are made.
The larval stage of some species, such as the brownish-yellow and black to
piceous Stripsipher zebra Gory & Percheron, is usually spent in rotten logs;
however, they are often found causing considerable damage to thatched roofs, as
in the case of the above-mentioned Trichostetha capensis. Some members of the
tribe Trichiini, such as Agenius limbatus (Olivier), are often found under com-
pletely dry cow-pats.
The tribe Cremastochilini contains several members which are either found in
birds’ nests, in ants’ and termite nests, or occur in beehives. At least two species,
Pseudospilophorus plagosus (Boheman), shiny black with two large whitish marks
on the elytra as well as an oblong whitish mark near the elytral apices, and about
12 mm long, and Brachymacroma emarginicollis (Boheman), of about the same
length and black with large yellowish patches, are predacious, feeding on the
yellow aphid, Aphis nerii Boyer de Fonscolombe, on milkweed plants (Asclepias
spp.) (Skaife 1953). Pseudospilophorus lugubris (Fabricius), similar to P. plagosus,
but lacking the distinct oblong mark on the elytral apices, often occurs in the nest of
the social spider (Stegodyphus sp.) of the family Eresidae.
According to Peringuey (1907) the species of Pseudospilophorus breed in the
nests of small birds such as finches, etc., feeding in both the larval and adult stage
on the faeces of the young birds. This also applies to the large, shiny, black hive-
beetle, Diplognatha gagates (Forster) (18-30 mm long—tribe Diplognathini),
the larvae of which construct a fairly hard earthen cocoon in which pupation
occurs, as is the case with the above-mentioned species.
Members of the genera Hoplostomus, such as the shiny black H. fuliginosus
(Olivier), a typical cetoniid (about 23 mm long), and Goniochilus, particularly
the black and dark-red G. bicolor Harold (about 22 mm long), are often attracted
to beehives where they feed on the honey. The shiny-black Genuchus hottentottus
(Fabricius), about 10 mm long, on the other hand, is commonly found in the
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
flowers of the sugar-bush (Protea repens), feeding on the nectar, and occasionally
appears in the carton nests of the cock-tail ant, Crematogaster peringueyi Emery,
in the Cape Peninsula. The reason for their presence in the ants’ nest is still
unknown, but from observations it seems certain that they are treated as per-
secuted synoeketes.
Species belonging to at least six genera of the tribe Cremastochilini have
been observed to be myrmecophilous or termitophilous, most of them apparently
being treated by the hosts as persecuted synoeketes as in the case of the species
belonging to the genus Genuchus. Coenochilus, the biggest genus of this particu-
lar section, includes about sixteen southern African species, most of which are
fairly widely distributed. Peringuey (1907) particularly mentions the large,
dark-brown C. hospes Peringuey (about 24 mm long) from the mounds of the
large fungus-grower termite, Odontotermes transvaalensis (Sj6stedt). Another
member, the dark-brown Trichoplus aegyptus Kolbe, often occurs in the mounds
-of the harvester termite Microhodotermes viator (Latreille) in Namaqualand.
Adults of about seven species were found to occur in the nests of the pug-
nacious ant, Anoplolepis custodiens (Smith), viz. Trichoplus vicinus Peringuey,
T. schaumi Westwood, Plagiochilus diversus Peringuey, P. intrusus Peringuey,
Scaptobius capensis (Gory & Percheron), Placodidus compransor Peringuey, and
Myrmecochilus marshalli Wasmann. They are all flattened, dark-brownish
beetles, about 9 mm long, except the last which is about 15 mm long and black
with a whitish band along the sides of the body. Species of the genus Trichoplus
mentioned above are easily recognized by their very concave prothorax.
Coenochilus appendiculatus Gerstaecker was apparently found in the nest of the
black sugar-ant, Acantholepis capensis Mayr, according to Schein (1954), as well
as in the mounds of a large fungus-grower termite (Macrotermes sp.).
The subfamilies Rutelinae and Melolonthinae include the chafer-beetles or
cockchafers and monkey-beetles. Both subfamilies are well represented in
southern Africa and contain species of economic importance.
In the Rutelinae, with about a hundred southern African species, the
adults have nine- to ten-segmented antennae and in the larvae the ninth and tenth
terga are distinct. In this case the anus is not angulate in the middle. At least
about seven species are harmful, including the maize-chafer beetle Chaetado-
retus cribrosus (Harold), the pecan-beetles Anomala probativa Peringuey and
Adoretus laticeps Fahraeus, the two wattle-chafers Anomala caffra Burmeister
and Adoretus ictericus Burmeister (Fig. 141), and sometimes also Adoretus tessu-
latus Burmeister, which feeds at night on rose leaves (Smit 1964). Adoretus
ictericus is often attracted to sites of decaying carcasses, mostly during the late
ceratophagous stage when the soil is enriched with organic matter; in this case
their larvae have on various occasions been recovered from the soil under the
carcasses.
Nearly 900 species of melolonthids occur in southern Africa, some of the
largest South African members being the dark-brown species, Macrophylla
SOUTH AFRICAN ARTHROPODS 259
pubens Peringuey (about 27 mm long) and M. maritima Burmeister (about
33 mm long), both from the southern and eastern Cape Province. The adults of
these beetles have nine- to ten-segmented antennae and in the larvae the ninth
and tenth terga are also distinct; the anus, however, is angulate in the middle in
this case.
The larvae of at least five species of Macrophylla cause severe damage to
golf- and bowling-greens in the eastern Cape Province. In Zimbabwe the large
brownish Eulepida mashona Arrow (about 25 mm long), with short decumbent
hairs giving it a greyish appearance, is regarded as a pest of maize, whereas the
smaller (about 15 mm long) shiny, dark-brown Schizonycha profuga Peringuey.,
of the same distribution, damages tobacco plants. According to the collection
data they do not occur in South Africa. Eucamenta castanea (Boheman)
(Fig. 14H), very similar to S. profuga and of about the same size and colour, is
sometimes attracted to decaying carcasses in the Cape Peninsula together with
Adoretus ictericus during the last post-mortem stage; its larvae, however, have
never been recovered under such circumstances.
The most abundant melolonthids are the monkey-beetles (tribe Hopliini),
which are mostly attracted to flowers, particularly burrowing in yellow and white
flowers of the family Compositae. One of the most common in the western Cape
is the hairy black Anisonyx ursus (Fabricius), which is about 10 mm long. These
beetles are usually very abundant for a short while during the spring, but then
disappear until the following spring. They are easily recognized by the enlarged
and long hind legs, especially of the males. Adults of most of the species are often
attracted to decaying matter. However, very little is known about their life-cycle:
it is probably the same as that of the small wattle-chafer, Monochelus calcaratus
Burmeister (see Prins 1965). Some of the largest members of this tribe are the
dark-brown Hoplocnemis hylax (Fabricius) and H. spectabilis Peringuey of the
eastern Cape Province, and H. koikoina Peringuey of Namaqualand (males of all
these species about 15 mm long).
Aphodius laetus Wiedemann
DESCRIPTION
Adult
Previously described by Peringuey (1901). Very similar to Aphodius procerus
(Fig. 16A) but smaller, being 6,6—8,3 mm long. Head and pronotum dark brown
to almost black; elytra lighter, yellowish brown to flavous; moderately shiny.
Widely distributed in the Cape Province; also found in Natal and Senegal
(Schmidt 1910).
Larva
Mature larva 9-10 mm long. Whitish to bluish white with semi-matt, brown
head; frons pale brown to almost yellowish and A. Jaetus is therefore easily
recognizable. Overall shape like that of A. procerus, but smaller. The thoracic
260 ANNALS OF THE SOUTH AFRICAN MUSEUM
spiracles oblong and somewhat larger than others; area in front of spiracle very
convex. Abdominal segments 1-6 with three annulets, segments 7—8 with two
annulets. Each annulet with a row of fine hairs; those on segments 2—5 short and
spine-like in older specimens. Legs sparsely covered with hairs, basal half of
coxae almost devoid of hairs. Claws simple and more distinctly demarcated than
in A. procerus (Fig. 15N).
Head (Fig. 151)
Epicranial suture distinct. Frons lighter coloured than rest of head, with four
anterior frontal and four posterior frontal setae, both forming two curved rows.
Mesal side of frons near base also with setae, as in A. procerus. Epicranium
darker than frons, as already stated, and with four to six lighter spots, each with a
long seta. Lateral sides of epicranium with numerous short setae. Antennae with
_ four large movable segments and a small fifth segment bearing sensilla. Setal
pattern very similar to that of A. procerus, except for very small setae on
epicranial sides.
Labrum and clypeus
Labrum slightly shorter than clypeus, the two clithra clearly visible dorsally.
Median row of four long setae present; posterior to this row are two shorter setae,
just in front of clypeal border. Median lobe dorsally with two long setae as well as
four strong setae on extreme anterior border as in A. procerus. One long and two
short setae also present laterad of each clithrum on lateral lobes. Clypeus about
1,25 times the length of the cranium, with one long and one short seta on each
side of postclypeus, as well as two shorter paramedian setae.
Epipharynx (Fig. 15K) very similar to that of A. procerus, but each acantho-
paria consisting of only four setae, of which one Is situated in front of each torma
as in latter species. Tormae fairly symmetrical and central area of pedium also
similar to that of A. procerus, but distal section of hairs flanking anterior part of
epitorma, more strongly developed than rest of hairs and much more pro-
nounced.
Mandibles (Fig. 15P)
Rather similar to those of A. procerus (Fig 16K), but neck of mandible in
region of scissorial teeth (t2 and t3) and molar area wider and only two setae
visible on exterior margin in specimens examined. Scrobis on each mandible
indistinct and dorsal carina rounded and weak. First molar tooth (m1) narrower
than in A. procerus and straighter; transverse groove on right molar area almost
obsolete; ventral section of molar area of right mandible forms separate piece
between m3 and ma, which is not so pronounced in A. procerus. Stridulatory area
absent on both mandibles. Dorsomolar setae (xy) on both mandibles similar to
those of latter species.
SOUTH AFRICAN ARTHROPODS 261
Fig. 15. Scarabaeidae. A—H. Aphodius moestus. A-D. Larva. A. Left lateral view. B. Apical
abdominal segment, ventral view. C. Head, dorsal view. D. Anal opening, viewed directly from
the rear. E. Adult. F. Egg. G—H. Larva. G. Epipharynx. H. Right mesothoracic leg.
I-P. Aphodius laetus. I-N. Larva. I. Head, dorsal view. J. Apical abdominal segment, ventral
view. K. Epipharynx. L. Left maxilla, dorsal view. M. Hypopharynx. N. Right mesothoracic
leg. O. Pupal hooks, ventral view. P. Mandibles, dorsal view (larva).
Q-S. Aphodius moestus larva. Q. Mandibles, dorsal view. R. Left maxilla, dorsal view.
S. Hypopharynx.
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
Maxillae (Fig. 15L)
Very similar to those of A. procerus; the galea free for most of its length.
Latter with a single tooth and five setae on dorsal side; ventrally with longitudinal
row of eleven to twelve strong setae, situated close together (as in A. procerus),
as well as single seta close to uncus. Lacinia ventrally with two setae of which
apical one is strong and tooth-like and situated near the three unci. Mala dorsally
with seven strong setae. Stipes with two dorsal setae and row of eight to eleven
small stridulatory teeth. Also two to three teeth near border of palpifer, which
has no teeth, but some papillae on its exterior border. Labacoria dorsally bare,
but with three setae on ventral side. Stipes also ventrally with one distal and one
proximal seta; latter located near border of galea. Palpifer and first segment of
maxillary palp each with long ventral seta; penultimate segment also with one
ventral and one external seta. Alacardo with one seta.
Labium
Very similar to that of A. procerus, distal sclerite of prementum as well as
proximal sclerite each with two long setae; postmentum with one or two setae on
each side; also two setae close together near anterior margin of distal sclerite of
prementum.
Hypopharynx (Fig. 15M) very similar to that of A. procerus; posterior tooth
of oncyli also strongly developed, but tooth t: very weak in specimens examined.
Ninth and tenth abdominal sterna (Fig. 15J)
Similar to A. procerus. Ninth sternite bearing transverse row of six setae;
division between the two segments clearly indicated. Raster: tegilla separate as in
A. procerus, each consisting of about twenty-five spine-like setae. Septum present
and wider posteriorly, flanked on each side by about ten to twelve pali. Otherwise
as in latter species. Anal lips similar to those of A. moestus, but both of the same
size in most specimens seen.
Pupa (Fig. 150)
Yellowish white and very similar to that of A. procerus, but smaller. Length
6,2—7,0 mm (posterior hooks excluded).
BIOLOGY
Aphodius laetus beetles were collected in fresh and semi-fresh cow-dung
almost throughout the year. Larvae were observed during the winter and spring
and were found together with those of A. consimilis. They were fairly heavily
parasitized by the dexiid, Pretoriamyia sellifera van Emden. When mature, the
larvae construct soft clay cells either in the soil or in the dung in which the
prepupae and pupae are formed.
The pupal stages lasted about 21 days in the laboratory during the winter and
12-14 days during the summer; adult beetles emerged during midwinter and early
summer.
SOUTH AFRICAN ARTHROPODS 263
Aphodius moestus Fabricius
DESCRIPTION
Adult (Fig. 15E)
Previously described by Peringuey (1901). Straw-coloured and fairly shiny,
particularly on pronotum; head and pronotum dark brown to almost black. Each
elytron with two narrow, longitudinal, blackish marks. On each side of pronotum
also brownish to blackish patch. Striae on elytra well indicated but not deeply
punctate. Length 5,8—7,5 mm.
Widely distributed in Subsaharan region, also present in Sri Lanka, Mada-
gascar and Malaysia (Schmidt 1910).
Larva (Fig. 15A)
Bluish white, with head dull, dark brown. Length 10,5—12,5 mm. Rather
similar to Aphodius laetus and A. procerus. Abdominal segments 1-5 each
divided into three annulets; segments 6-8 apparently with two annulets each.
Sparsely covered with setae all over the body, with most setae on abdominal
segments 1—5 small and spine-like. Each abdominal sternum with transverse row
of four to six setae on middle. All legs more or less of same size, well developed
and sparsely covered with setae, particularly coxae; claws (Fig. 15H) simple and
acute, lacking two spine-like setae present in A. /aetus; ventral surfaces of claws
serrate and studded with minute denticles. Both femur and tibia with minute
denticles on ventral side (absent in A. /aetus).
Head (Fig. 15C)
Frontal sutures rather indistinct in old specimens, epicranial stem (coronal
suture) as in A. /aetus. Frons somewhat lighter in colour than epicranium in some
specimens, in others just as dark and bearing only three long setae on each side.
Two short paramedian setae or spines also present below frontoclypeal suture,
forming part of anterior frontal setae. Epicranium with anterior corners darker in
most specimens seen, dorsally with four setae as in A. /aetus, as well as some
minute setae on lateral margins, which are, however, not as conspicuous as in
latter species. Antennae slightly shorter than cranium, with four large and small
fifth segments; separation of the first two segments very indistinct.
Labrum and clypeus
Setal pattern of labrum as in A. Jaetus, except two posterior setae immedi-
ately in front of clypeal border absent. Clypeus about as long as labrum; very
convex in lateral view and with one long seta on each side, as well as two smaller
paramedian setae.
Epipharynx (Fig. 15G) as in A. laetus, except crepis is much better indicated
and distal section of hairs flanking anterior part of epitorma larger, covering
almost whole of protophoba. Two or three strong setae in central part of dexio-
phoba are characteristic of both this species and A. /aetus; they are not as obvious
in A. procerus.
264 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mandibles (Fig. 15Q)
Stridulatory areas absent in both mandibles. Lateral side of each mandible
rounded dorsoventrally, without distinct dorsal carina and with imperfect scrobis.
Only two small setae observed on each mandible of which proximal one is
situated on lateral face. Cutting edges very similar to those of A. laetus; apical
tooth much longer on both mandibles. On left mandible scissorial notch much
more distinct and second tooth (ti) fairly obvious in most specimens, giving
cutting edge a tridentate appearance. In both mandibles the neck much longer
than in A. /aetus. Molar areas of both mandibles similar to those of the latter;
dorsomolar setae, however, arranged in form of transverse row on left mandible.
Maxillae (Fig. 15R)
Similar to that of A. /aetus. Galea and lacinia free apically, latter with seven
strong setae on dorsal side and apically with three unci. Ventrally with two setae
only, apical one spine-like as in A. laetus. Galea bears single uncus, with two
adjacent curved setae as well as longitudinal row of closely adjoining setae
ventrally; dorsally with about seven strong setae of which two are situated on
lateral side of uncus. Stipes with two setae dorsally and a row of eight to nine
stridulatory teeth; also about three stridulatory teeth near base of palpifer;
ventrally with only single proximal and single distal seta, as in A. /aetus. Palpifer
and basal palpal segment with single ventral seta; penultimate segment with one
ventral and one lateral seta. Labacoria with three ventral setae.
Labium
Similar to that of A. /aetus, but with only two long setae on postmentum.
Hypopharynx (Fig. 15S) also very similar to that of A. /aetus, except that
hypopharyngeal sclerite is somewhat better sclerotized. Glossa with same number
of setae as in latter species.
Ninth and tenth abdominal sterna (Fig. 15B)
Distinct line of demarcation between ninth and tenth sternum present as in
A. procerus. Raster very different from that of either A. procerus or A. laetus, as
septum is absent. Two tegilla more or less fused, and with about seventy small
black tubercles, each with small hamate seta (almost as in Oniticellus). Barbula
absent. Anal opening visible as transverse slit (Fig. 15D), lower anal lip smaller
than upper lip and divided ventrally by acute emargination; lateral margin on
each side slightly emarginate in most specimens seen; very similar to A. /aetus and
A. procerus, but ventral incision much deeper.
Pupa
Similar to that of A. procerus (Fig. 16L), including caudal hooks. Length
6,5—7,4 mm (excluding caudal hooks). Colour pale whitish brown, but turning to
dark colour a few days before beetle emerges.
SOUTH AFRICAN ARTHROPODS 265
BIOLOGY
Eggs of Aphodius moestus (Fig. 15F) were found in fresh cow-dung near
Montagu during the summer months. They are whitish, oval and matt, 1,5 mm
long and 1,0 mm broad. All the eggs collected during the summer had an
incubation period of at least 5—6 days, the young grubs being about 2,5 mm long
just after hatching. At first they are almost pure white, but soon the head
becomes darker, almost golden brown. The setal pattern of the first instar is
almost the same as that of the mature larva, except that the very small para-
median setae on the clypeus and the median pair of anterior frontal setae are
absent. In all specimens seen, the posterior pair of dorso-epicranial setae (desz) is
also absent.
Large numbers of full-grown larvae and pupae were found in fresh dung
during the autumn and beetles were observed almost throughout the year. The
life-cycle from egg to adult during December and January occupied 30-33 days.
All the pupae collected were found in small clay cells in the partly dry dung which
remained after the feeding of the various coprophagous insects.
The newly-emerged beetle is pale brownish yellow, almost straw coloured on
the elytra as well as on the pronotum, with the central part of the latter brownish;
the head is brown and the legs yellowish. After a day or two it assumes its normal
colour.
Aphodius procerus Harold
DESCRIPTION
Adult (Fig. 16A)
Previously described by Peringuey (1901). Brownish in colour, with
prothorax darker in middle and usually finely pitted. Elytra striate, the striae with
one row of punctures each. Head also finely punctate. Prothorax about as wide as
elytra and whole insect moderately shiny. Length 11—12,5 mm.
Widely distributed in the western Cape (Peringuey 1901).
Larva (Fig. 16H)
Mature larva measures 16-17 mm in length. Colour bluish white, with head
dark blackish brown to almost black, and matt. Abdominal segments 1-6 with
three annulets; setae on segments 2-6 short and almost spine-like. Legs sparsely
covered with setae. Claws (Fig. 16G) simple and acute, each bearing two small
setae near base. Each abdominal sternite, except segment 10, with transverse row
of six to eight long setae.
Head (Fig. 161)
Frontal sutures distinct. Areas around antennal bases whitish, anteclypeus
and labrum yellowish to brownish yellow. Frontoclypeal suture distinct. Frons
with only two long setae on each side and also two short paramedian anterior
frontals. Epicranium with four setae on dorsal side as in Aphodius laetus and
266 ANNALS OF THE SOUTH AFRICAN MUSEUM
anterior transverse row
a“
we
AIAN
Fig. 16. Scarabaeidae. Aphodius procerus. A. Adult. B-K. Larva. B. Anterior margin of
labrum, viewed directly from the front. C. Epipharynx. D. Left maxilla, dorsal view.
E. Hypopharynx. F. Left maxilla, ventral view. G. Right mesothoracic leg. H. Larva, left
lateral view. I. Head, dorsal view. J. Apical abdominal segments, ventral view. K. Mandibles,
dorsal view. L. Pupa, ventral view.
SOUTH AFRICAN ARTHROPODS 267
A. moestus. Antennae slightly more than half the length of cranium, with five
movable segments of which apical one is very small and bears two oval sensory
grooves.
Labrum and clypeus
Labrum divided by clithra; median lobe bearing four strong setae dorsally on
extreme anterior margin and two long setae further posteriorly (Fig. 16B). One
long and two short setae (x) present laterad of each clithrum on lateral lobes
(Fig. 16B, I); posterior to these are two transverse rows of four setae each; in
some specimens posterior row contains only two to three setae. Clypeus of about
same length as labrum and bearing transverse row of four setae on postclypeus.
Epipharynx (Fig. 16B-—C). Distinct clithrum present on each side of corypha;
latter with four medium long spine-like setae. Pedium large, raised and encircled
by setae of proto-, dexio- and laeophobae. Tormae (dt and It) narrow and merely
indicated by suture; anterior epitorma elongate and ending in central cavity. Each
torma with short spine on lateral margin, which apparently forms part of acantho-
paria. Central area of pedium concave; anterior epitorma flanked on right by fine
hairs. Each acanthoparia with five short stout spines and distally with two longer
spine-like setae. Chaetoparia and acroparia absent. Boundary line of epipharynx
and pharynx not clearly indicated by crepis in specimens examined; area enclosed
by latter (also known as haptolachus, which is actually situated on clypeus) only
with triangular sclerotized plate, in front of which is transverse row of fine hairs.
Mandibles (Fig. 16K)
Nearly as long as the cranium; dorsal carinae rounded and not clearly
indicated; scrobis imperfect and bearing three short, sickle-shaped setae. Cutting
edge of left mandible blade-like; scissorial notch distinct on both mandibles.
Posterior scissorial tooth (ts) on the right mandible rounded in most of specimens
examined. Molar area of left mandible divided into distal and proximal lobe;
distal lobe (m:) with broad transverse cutting edge, with curved row of dorso-
molar setae (xy); proximal lobe (m2) broad, its dorsal aspect sharp. Molar area of
right mandible entire, with transverse groove in front of small ventral tooth (ms)
and bearing fine setae (xy); proximal tooth (mz) pointed posteriorly. Fine setae on
posterior part of molar area of both mandibles present as brustia.
Maxillae (Fig. 16D, F)
Galea and lacinia fused, but delimited dorsally and ventrally by deep suture;
apices free. Galea bearing single strong apical uncus and two to three strong
curved setae; dorsally with longitudinal row of about five strong setae and
ventrally with another row of closely approximated slender setae. Lacinia with
three well-developed apical unci. Mala dorsally with row of nine to eleven strong
setae, and ventrally with only one strong seta near unci; also with another seta
close to its base. Stipes dorsally with two setae close to base of palpifer as well as
well-defined carina (c) with row of about nine stridulatory teeth. Base of palpifer
268 ANNALS OF THE SOUTH AFRICAN MUSEUM
also with four stridulatory teeth; ventrally with single seta. Labacoria bare on
dorsal side but ventrally with two long setae. Alacardo on both sides with single
seta. Maxillary palp with four movable segments; ventral side of basal segment
with single seta and also one on both external and ventral sides of penultimate
segment.
Labium
Mentum trapezoidal, with strong seta on each side near base. Proximal
sclerite of prementum wider than distal sclerite, fairly convex transversely in
lateral view and with two paramedian setae. Distal sclerite with two setae close
together near anterior margin; sclerite consisting almost of two parts due to
presence of shallow longitudinal depression; each part with long median seta.
Hypopharynx (Fig. 16E). Distal region of glossa with three setae on each
side. Transverse ridge in front of oncyli bears row of fine setae. Hypopharyngeal
- sclerite concave in middle; oncyli consisting of at least three strongly sclerotized
teeth (ti, t2, t3). Lateral sclerites not strongly developed. Oblique row of fine setae
present on each side, laterad of oncyli, each row continuing anteriorly over lateral
area of glossa, terminating on ventrolateral side of distal sclerite of prementum in
three strong setae.
Ninth and tenth abdominal sterna (Fig. 16J)
Dividing line between these segments distinct. Lower anal lip (lamina sub-
analis) divided into two sections in middle; upper anal lip (lamina supra analis)
entire; both lips demarcated by thin, well-defined suture. Anus present as trans-
verse slit. Barbula absent. Raster: tegilla separate; each tegillum with eleven to
thirteen strong setae similar to pali, as well as some smaller spine-like setae.
Septum long, wider in the middle; each pallidium with fourteen strong pali.
Campus small, bearing only one long seta on either side of tegilla. Oblique row of
long setae also present in front of anal lips.
Pupa (Fig. 16L)
Light or pale brownish, 11-12 mm long. Bare, almost without setae. Abdo-
men dorsomedially with carina as in A. /aetus. Wing-covers longitudinally striate.
Extreme apex of abdomen carrying two long hooks, which are curved inwards at
tips. Few days before beetle emerges, pupa turns dark brown to piceous.
BIOLOGY
Aphodius procerus beetles were mainly observed during the winter and
spring. The larvae feed on semi-fresh to semi-dry cow-dung and were collected
from late autumn to early spring; they are easily recognized by the dull, dark-
brown to almost black heads, which are white after each moult. Before the
prepupal stage is reached the larva constructs an earthen cell either in the dung or
in the upper few centimetres of soil, in which the prepupal and later, after a few
SOUTH AFRICAN ARTHROPODS 269
days, the pupal stage is formed. Pupae were found during the winter and early
spring. Pupal stages lasted from 17—23 days and beetles emerged from August to
October.
Aphodius consimilis Boheman
DESCRIPTION
Adult
Previously described by Peringuey (1901). Rather similar to, but somewhat
shinier than Aphodius moestus and without longitudinal black stripes on elytra;
punctato-striations less distinct. Smaller than latter, only 4,2—5 mm long.
Widely distributed in the Cape Province, also present in Mozambique
(Schmidt 1910).
Larva
Very similar to A. moestus. First six abdominal segments each with three
annulets, seventh and eighth apparently with only two. Setal pattern similar to
A. laetus, but lacking denticles present in A. moestus. Smaller than those of
above two species, measuring only 7-8 mm when mature. Body bluish white;
head shiny golden brown. Each abdominal sternum with about eight (or slightly
more) setae.
Head (Fig. 17C)
About 1,25 times wider than long. Coronal suture as in A. /aetus and
A. moestus. Frons with transverse row of four setae near frontoclypeal suture,
which is well defined; also with long seta on each side, laterad of antennal base.
Epicranium with longitudinal row of three dorso-epicranial setae as well as one
exterior dorso-epicranial seta on each side; anteriorly with single long seta near
antennal base. Antennae slightly more than half the length of cranium, appearing
four-segmented; the first and second segments almost immovably united; apical
segment small, bearing at least one large sensory spot and some sensory pegs;
penultimate segment also with sensory grooves on protuberance as in other
species.
Labrum and clypeus
Two well-defined clithra present, dividing labrum into small median and two
lateral lobes. Setal pattern of labrum as in A. /aetus; also with two smaller setae
near clypeal border. Clypeus slightly longer than the labrum, with same setal
pattern as in latter species.
Epipharynx (Fig. 17E) very similar to that of A. laetus with same setal
pattern; also with one or two strong setae in middle of dexiophoba. Tormae fairly
symmetrically developed and crepis fairly distinct.
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 17. Scarabaeidae. A-—G. Aphodius consimilis larva. A. Apical abdominal segment, ventral
view. B. Anal opening, viewed directly from the rear. C. Head, dorsal view. D. Mandibles,
dorsal view. E. Epipharynx. F. Left maxilla, dorsal view. G. Hypopharynx.
H-M. Onitis aygulus. H-L. Larva. H. Epipharynx. I. Labrum and clypeus, dorsal view.
J. Hypopharynx. K. Left maxilla, dorsal view. L. Mandibles, dorsal view. M. Egg.
SOUTH AFRICAN ARTHROPODS Jogi
Mandibles (Fig. 17D)
About 1,2 times wider than long. Rather similar to those of A. Jaetus
(Fig. 15P), with neck very short and the lateral sides rounded. Dorsal carina
rounded and scrobis indistinct. Three setae visible on dorsal side, of which one is
situated on lateral face. Cutting edge of left mandible broad, scissorial notch
clearly indicated. Second tooth (t:) fairly blunt or rounded. Cutting edge of right
mandible similar to that of A. procerus, but second tooth (t3) broad and in
some specimens almost bilobed. Molar areas similar to those of A. laetus and
A. procerus (including dorsomolar setae). Both mandibles with brustia.
Maxillae (Fig. 17F)
Galea and lacinia free apically as in other Aphodius species; galea with a
single uncus and dorsally with five setae. Ventrally galea bears longitudinal row of
closely approximated setae as in A. procerus as well as a single seta on lateral
margin near apex. Lacinia with three well-developed unci, of which distal one is
largest, and single ventral seta, situated close to proximal uncus. Dorsally there
are six strong setae near mesal margin. Stipes dorsally with row of about eight
stridulatory teeth of which at least five are fairly conspicuous; two or more
stridulatory teeth and single seta located near base of palpifer. In the other
Aphodius species described here there are two setae in this position. Ventrally
there are two setae as in A. /aetus; four-segmented maxillary palp, including its
setae, as in A. /aetus. Labacoria dorsally bare, ventrally with two setae. Alacardo
with a single seta.
Labium
Similar to that of A. /aetus and A. moestus, but the postmentum with only
one seta on each side.
Hypopharynx (Fig. 17G) similar to the other three species described above.
Hypopharyngeal sclerite well developed, oncyli almost as in A. procerus. Lateral
sclerites fairly well developed; transverse row of setae in front of the
hypopharyngeal sclerite indistinct.
Ninth and tenth abdominal sterna (Fig. 17A)
Rather similar to those of A. moestus, the two segments clearly demarcated.
Raster consisting of two tegilla, which are fused proximally but separated distally
in some specimens. Each tegillum with twenty-three to twenty-five hamate setae;
in old specimens each seta situated on tiny black tubercle as in A. moestus.
Barbula absent. Anal lips (Fig. 17B) similar to A. moestus, but lower anal lip
much larger and in most specimens slightly larger than upper anal lip.
Pupa
Similar to that of the other Aphodius species described here (including
abdominal hooks), but smaller. Length 4,2—4,9 mm. Yellowish white in colour
but becoming dark brown just before emergence of imago.
DAD ANNALS OF THE SOUTH AFRICAN MUSEUM
BIOLOGY
Aphodius consimilis is one of the most numerous scarabs found in fresh to
semi-dry cow-dung in the southern and western Cape. It is usually associated with
other species, such as A. moestus, A. discoidalis, A. procerus, etc. Larvae of this
species were collected almost throughout the year, except in midwinter. The eggs
resemble those of A. moestus and measure from 0,56 by 0,44 mm to 1,0 by
0,64 mm. They are slightly shiny, dirty white and almost transparent and, as in all
other scarab eggs, they increase in size after oviposition. All eggs were laid in
semi-fresh to semi-dry cow-pats, in groups of five or more, and were never found
in fresh dung. When mature, the larva constructs a little clay cell in which the
whitish prepupa and later the pupa is formed. If dung is still available, the cells
are constructed in the remains of the pats, otherwise in the top layer of the soil.
According to observations made the incubation period of the eggs seems to be the
same as for A. moestus. The larval period occupied 19-25 days during August to
September and the pupal stage 16—18 days. The whole life-cycle was completed in
35-47 days in specimens kept in the laboratory (16-22 °C). Newly-emerged
beetles are straw coloured.
Onitis aygulus (Fabricius)
DESCRIPTION 3
Adult
Described by Peringuey (1901) and Skaife (1953). Thorax bronzy green,
elytra more greenish brown. Similar to Onitis caffer in general form and shape.
Elytra with seven distinct striae. Length varying from 18,3 to 23 mm.
Widely distributed in the western and north-western Cape; also present in
South West Africa (Ferreira 1978).
Larva
Larva rather similar to that of O. caffer, as described by Oberholzer (1958).
As only few mature specimens were available for study, it was difficult to deter-
mine exact number of annulets, but abdominal segments 1—6 appear to have three
annulets and segments 7 and 8 only two. All annulets, except probably first two,
with scattered setae and those on at least first three segments arranged in single
transverse rows; spine-like setae occur on segments 3—10. Spiracles and legs as in
O. caffer. Sterna almost devoid of setae, except tenth sternum.
Head
Similar to that of O. caffer. Fairly smooth and colour golden, yellow brown,
marbled with iridescent pale blue on certain areas in some specimens. Ocelli
absent. Antennae four-segmented; apical segment small and bearing sensory
cones. As very few specimens were available, a reconstruction of the mouth-parts
SOUTH AFRICAN ARTHROPODS 273
was made from cast skins and compared with those that could be preserved for
drawing.
Labrum and clypeus (Fig. 171)
Differ from those of other scarab larvae described here by presence of large
numbers of setae, particularly on labrum. Two clithra divide latter into three
rounded lobes, each bearing large number of setae, which are also continuous on
median area. Postclypeus bearing median transverse row of nine to ten long
setae. Both the clypeus and labrum seem to agree well with those of O. caffer, but
judging by the figures of Oberholzer (1958), labrum more setose in latter.
Epipharynx (Fig. 17H) rather similar to that of O. caffer, but acanthopariae
seem to be represented by only two or three setae on each side. Each chaetoparia
with at least twenty-two to twenty-six setae; acroparia with about six setae each;
corypha with about twelve strong rounded spines. Posterior epitorma better
developed than anterior one and two sensilla present anterior to protophoba.
Mesophoba sclerotized on left side and forms part of laeotorma; lateral lobes
each with small strongly sclerotized area. Fused tormae, and anterior and pos-
terior epitormae fairly well developed. Otherwise as in O. caffer.
Mandibles (Fig. 17L)
Reddish brown, cutting and molar areas black. Only slightly longer than
wide and nearly as long as cranium. Dorsal carinae rounded, scrobis indicated by
short shallow depression with about four setae; also single curved seta present in
middle of neck between teeth t: and tz. Stridulatory areas absent on both man-
dibles. Judging by Oberholzer’s figures, mandibles are fairly different from those
of O. caffer. In O. aygulus, cutting edge of left mandible is clearly tridentate;
scissorial notch fairly wide and distinct. On right mandible cutting edge is
bidentate with wide and deep notch between first and second tooth (ts). Molar
area of left mandible clearly divided into proximal (mz) and distal (m1) lobe; latter
long and broad. On right mandible division not so clear, molar area forming a
wide tooth (m3) ventrally. Short oblique row of fine dorsomolar setae (xy) present
on each mandible.
Maxillae (Fig. 17K)
Rather similar to those of O. caffer. Galea and lacinia free, latter with single
strong uncus, its mesal margin beset with strong setae. Ventral side of lacinia with
numerous setae. Galea with very small uncus surrounded by curved row of about
eight rounded spines, which continue as strong setae on ventral side. Whole of
ventral area studded with hamate setae, its dorsal surface with numerous strong
setae. Palpifer of three-segmented maxillary palp with about fourteen setae.
Stipes with one long and some short ventral setae; dorsally with about eighteen
setae and sixteen conical stridulatory teeth. Penultimate segment of maxillary
palp with two setae on lateral margin.
274 ANNALS OF THE SOUTH AFRICAN MUSEUM
Labium
Similar to that of O. caffer, palpi with apical segment fairly long.
Hypopharynx (Fig. 17J). Glossa with about ten hairs in middle and numer-
ous setae on right side. Left side with only two setae in specimens examined.
Right lateral sclerite with numerous spine-like setae; left one with only twenty to
twenty-two spine-like setae. Oncyli on right side with two well-developed teeth
(t: and tz); third tooth (ts) more weakly developed.
Ninth and tenth abdominal sterna (Fig. 18C)
Rather similar to those of O. caffer. The two sterna clearly demarcated. The
two palidia long, each with eighty-seven to ninety-two pali, anterior third sep-
arated from the rest. Tegilla as in O. caffer, each one with numerous palus-like
setae. Septum long, somewhat wider posteriorly. Anal lips similar to those of the
latter species.
BIOLOGY
Skaife (1953) describes the biology of Onitis aygulus and according to him
the beetle first digs a tunnel in the soil beneath the dung. At the bottom of this
tunnel a large chamber is excavated into which a mass of dung is carried and in
this mass the eggs are deposited at intervals as the filling of the chamber proceeds.
The cells in which the eggs are laid are always made near the surface of the dung.
About six white, oval eggs (Fig. 17M) are laid and these hatch within about
14 days. When fully grown each larva makes a cell in which pupation occurs. The
pupal stage lasts 14—21 days.
In sandy areas along the west coast, the oblong whitish eggs of this species
were often observed in semi-fresh cow-pats during January. Fairly large larvae
(about 35 mm long) that hatched from such eggs were collected from hollowed-
out chambers in the pats in the same areas during the end of January and
beginning of February. Pupae from these larvae were about 16 mm long, pale
lemon yellow, and remained in the cells for about 38 days during September
before the beetles emerged. Under such conditions larval stages lasted for about
202 days before pupation occurred and the newly-emerged beetles were reddish
brown on the thorax with yellowish elytra and reddish legs.
Oniticellus pictus (Hausmann)
DESCRIPTION
Adult (Fig. 18A)
Previously described by Peringuey (1901). Length 6,4—8,7 mm. Pale yellow-
ish or flavous with large black triangle on head, large black patch on pronotum
and some black spots on elytra. Pronotum broader than elytra with conspicuous
pits which are usually obsolete on central part; also small black spot on each side
of pronotum. Legs flavous, marked with black.
SOUTH AFRICAN ARTHROPODS 275
Fig. 18. Scarabaeidae. A. Oniticellus pictus adult. B. Oniticellus pictus pupa, dorsal view.
C. Onitis aygulus larva, apical abdominal segments, ventral view. D-—K. Oniticellus pictus.
D-I. Larva. D. Head, dorsal view. E. Apical abdominal segments, ventral view.
F. Epipharynx. G. Left maxilla, dorsal view. H. Hypopharynx. I. Mandibles, dorsal view.
J. Pupa, ventral view. K. Apex of pupa, left lateral view. L. Oniticellus planatus, apex of pupa,
left lateral view.
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
Attracted to fresh and semi-fresh cow-dung as in the case of Oniticellus
planatus. Widely distributed in South Africa; also present in Zaire and east Africa
(Janssens 1953).
Larva
Very similar to that of O. planatus (Fig. 19C), but smaller. Abdominal
segments 5—8 with two rows of dorsal setae; fourth segment dorsally with short
spines only. Prothoracic shield angular on each side, but not toothed.
Head (Fig. 18D)
Only slightly wider than long, similar to that of O. planatus and having same
depressions. Frons with transverse row of four setae near clypeofrontal suture
and posterior to this two smaller paramedian setae; also single seta present on
each side near middle of frontal suture. Single seta present just mesad of each
~ antennal base. Epicranium anteriorly with three setae and posteriorly with longi-
tudinal row of four to five setae as in O. planatus. Sides of epicranium with very
small setae.
Labrum and clypeus
Labrum about twice as wide as long with the same number of setae as in
O. planatus. Clypeus also twice as wide as long, about one-third longer than
labrum and with two long paramedian setae and two lateral setae on each
side.
Epipharynx (Fig. 18F) very similar to that of O. planatus but clithra not
sclerotized. In mature larvae corypha and lateral lobes strongly sclerotized, as
well as anterior and posterior epitormae. In all specimens examined each acro-
paria with one seta. Mesophoba in most specimens seen with long setae. At least
one pair of macrosensilla clearly visible in most specimens.
Mandibles (Fig. 181)
Slightly more than half the length of cranium. Dorsal carina rounded and
scrobis indistinct and, as in O. planatus, with only two setae dorsally. Cutting
edge of left mandible with three teeth, scissorial notch distinct. Molar area
divided into two lobes; distal one (mi) long, broad and emarginate in middle;
proximal lobe (mz) as in O. planatus, with a fringe of hairs posteriorly. Right
mandible similar to that of O. planatus, but molar area with strong ridge (ms)
anterior to it. Each mandible also with longitudinal row with fine dorsomolar
setae.
Maxillae (Fig. 18G)
Similar to that of O. planatus. Galea with single short uncus, dorsally with
seven setae, ventrally beset with about twenty-five setae on mesal area. Lacinia
with single long uncus and eight strong setae dorsally; ventrally with five to six
setae of which the apical one is very short and spine-like in young specimens.
SOUTH AFRICAN ARTHROPODS OFT
Labacoria ventrally with two small setae; dorsally bare. Alacardo with about
three setae. Stipes ventrally with two setae and dorsally with three of which
posterior one is longest. Three inconspicuous stridulatory teeth also present.
Palpifer with two setae situated close together ventrally; penultimate segment of
maxillary palp with single long seta on the ventral side and one on lateral margin.
Labium
Postmentum trapezoidal; in mature specimens anterior part with broad
rounded ridge bearing only two short setae near base. Proximal sclerite of
prementum convex, transversely divided by shallow notch into two halves, each
with three setae, inner ones longest. Distal sclerite of prementum divided by
depression into two globular lobes, each with ten to twelve setae.
Hypopharynx (Fig. 18H). Glossa deeply cleft by depression into two halves,
each with five to seven setae; oncyli forming two strong teeth on right side, as in
O. planatus; left one with transverse tooth. Oncyli anteriorly encircled by a row of
strong setae in front of which there is transverse row of six setae. Each lateral
sclerite with row of fine hairs laterad of oncyli.
Ninth and tenth abdominal sterna (Fig. 18E)
Ninth and tenth segments clearly demarcated. Raster similar to that of
O. planatus, but iridescent areas oblong and narrow, beset on each side with
twenty to thirty tiny black tubercles, each with tiny hamate seta. Tenth sternite in
form of wide V-shaped lip just below lower anal lip (in O. planatus it is widely
convex, not V-shaped); otherwise as in the latter species. Barbula absent. Lateral
areas of tenth segment beset with small setae.
Pupa (Fig. 18B, J-K)
Almost pure white when formed, but turning darker as imago develops.
Length about 10 mm and, as in O. planatus, prothorax with large convex median
raised area and inconspicuous swelling on each pronotal shoulder. Abdomen with
four lateral projections, as well as pteronotal, dorsal tergal, and caudal support
projections (ptp, tp and cpp). Extreme apical part of dorsal projection, as well as
caudal projection and developing genital capsule fairly rounded so that there
appear to be three rounded knobs on abdominal apex (in O. planatus there are
only two which are fairly rounded, extreme apical part of caudal support projec-
tion not rounded but carinate).
BIOLOGY
As in Oniticellus planatus, the full-grown larvae of O. pictus are found in oval
clay cells somewhat smaller than those of the former species, i.e. 10-11 mm in
diameter. Prepupae and pupae are almost pure white and were collected during
March and April together with those of O. planatus in the same cow-pats. Beetles
emerged during April. Both these species seem to play an important part in the
destruction of the dung, as large numbers of pupal cells were collected per
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
cow-pat in certain areas around Mossel Bay and in the Brandwag area between
Mossel Bay and Oudtshoorn.
Oniticellus planatus Castelnau
DESCRIPTION
Adult (Fig. 19A)
Previously described by Peringuey (1901). Shiny black, elytra longitudinally
striate, each stria with one row of punctures. Prothorax somewhat wider than
elytra. Length 9,9-10,9 mm. Attracted to fresh and semi-fresh cow-dung and
very numerous in the Mossel Bay area during the surveys. Widely distributed in
Africa (Peringuey 1901; Janssens 1953).
Larva (Fig. 19C)
White in colour. Only pro- and mesothoracic segments with dorsal setae;
metathoracic segment almost devoid of setae. Abdominal segments 3-7 with
scattered setae dorsally; segments 8—9 apparently with two rows of setae; second
abdominal segment with one row of setae dorsally. Legs with some setae, mostly
on ventral side, coxae almost devoid of hairs; claws absent (Fig. 19H), but there is
short terminal seta surrounded by circle of six to eight hairs. Prothoracic shield
with transverse ridge on each side which is somewhat pointed in middle, but
lacking triangular projections as in Onitis caffer (Oberholzer 1958). Spiracles as in
O. caffer and with fairly long seta posterior to each spiracle.
Head (Fig. 19E)
Only slightly wider than long, frontal suture forming an inverted M, with
arms not very clearly indicated anteriorly. Coronal suture extended to middle of
cranium where it joins V-shaped notch or depression. Depression on fronto-
clypeal suture extending on to posterior half of postclypeus. Frons with three long
setae on each side as illustrated, two of which are situated medially, one on either
side of depression near clypeal border; another pit present exterior to each of
these setae. Epicranium with seven to eight dorsal setae on each side of which
four are situated in longitudinal row near the frontal suture on a narrow area that
is lighter in colour than rest of the cranium.
Labrum and clypeus
Labrum slightly less than twice as wide as long and deeply emarginate on
each side of corypha; lateral lobes rounded. Middle lobe with about seven to eight
strong setae; lateral lobes with four strong setae, the posterior two sickle-shaped;
there are also two paramedian setae. Clypeus only slightly longer than labrum,
with two lateral setae on each side and two paramedian setae on postclypeus.
Epipharynx (Fig. 19J). Clithra well developed and corypha with four strong,
rounded spines. Each chaetoparia with eight setae and each acroparia represen-
ted by two strong setae. Pedium raised above level of lateral lobes and encircled
SOUTH AFRICAN ARTHROPODS 279
Fig. 19. Scarabaeidae. Oniticellus planatus. A. Adult. B. Clay cell. C—K. Larva. C. Left
lateral view. D. Apical abdominal segments, ventral view. E. Head, dorsal view. F. Left
maxilla, dorsal view. G. Hypopharynx. H. Right mesothoracic leg. I. Anal opening, viewed
directly from the rear. J. Epipharynx. K. Mandibles, dorsal view.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
by dexio-, laeo- and protophobae. Tormae (dt and It) present as narrow sclerite;
both anterior and posterior epitormae present. Crepis represented by three
weakly sclerotized areas. Anterior half of area enclosed by crepis (haptolachus)
raised; upper surface almost flat and surrounded by mesophoba.
Mandibles (Fig. 19K)
Slightly more than half the length of cranium; dorsal carina rounded; only
two dorsal setae present and single ventral seta near molar area. Scrobis indistinct
and stridulatory area absent. Distal cutting edge of left mandible narrow, scis-
sorial notch in form of wide depression between apical tooth and molar area; also
single small tooth (t:) present below apical tooth. Molar area bilobed, distal tooth
(mi) broad and fairly sharp; proximal lobe (m2) with arcuate posterior margin,
bearing setae. Right mandible with two fairly rounded apical teeth, scissorial
notch shallow. Molar area consisting of single broad oblique lobe that forms fairly
~ strong tooth (mz) pointing backwards; ventral tooth separated only at posterior
-end. Molar area on each mandible with small groove on dorsal side bearing fine
dorsomolar setae.
Maxillae (Fig. 19F)
Galea and stipes free, each with single short uncus. Galea on dorsal side with
about seven setae, and ventrally with twenty to thirty long setae on mesal side.
Lacinia dorsally with about eight strong setae and ventrally with about five. Stipes
dorsally with four setae as well as row of about eight small stridulatory teeth.
Palpifer with two exterior setae and one long ventral seta. Penultimate segment of
maxillary palp with single exterior and single ventral seta. Labacoria bare, except
for two small ventral setae; alacardo with about four setae.
Labium
Distal sclerite of prementum divided by a depression into two almost conical
halves, each bearing ten to twelve setae. Proximal sclerite of prementum convex
and bearing median transverse row of six setae. Postmentum trapezoidal with two
short setae near base.
Hypopharynx (Fig. 19G). Glossa widely emarginate apically, each half bear-
ing about eight strong setae; hypopharyngeal sclerite strongly sclerotized, oncyli
forming at least two strong conical teeth (t: and t2). That part of glossa in front of
oncyli strongly sclerotized, forming ridge bearing fine hairs. Anterior to this
sclerotized area there is transverse row of about five short spine-like hairs.
Longitudinal row of four spine-like setae present on each lateral side of
oncyli.
Ninth and tenth abdominal sterna (Fig. 19D)
Ninth and tenth segments clearly divided, with inconspicuous, longitudinal
ridge on tenth sternite, ending in front of lower anal lip in small V-shaped notch.
SOUTH AFRICAN ARTHROPODS 281
On either side of this ridge an almost oval iridescent bluish patch present and
posterior to this a small tegillum bearing about twenty short, palus-like setae.
Single long setae situated laterad of each iridescent patch, as well as some smaller
scattered hairs. Anal opening transverse, surrounded by large upper anal lip
(Fig. 191) and smaller lower lip, divided into exterior and interior lobe. Barbula
absent.
Pupa
Very similar to that of O. pictus and when newly formed almost pure white,
but as imago develops it turns darker. Most of pupae collected range in length
from 12 to 13 mm. Extreme apex of abdomen with two swellings, the dorsal one
being the caudal support projection and the lower one the developing genital
capsule (Fig. 18L). Longitudinal ridge or dorsal tergal support projection more or
less carinate and obsolete on segments 4-7.
BIOLOGY
The creamy-white, almost pear-shaped eggs of Oniticellus planatus are laid
singly in a small dung ball, 14,5—15 mm diameter, in which the larva feeds. When
fully grown a clay cell (Fig. 19B) of similar size and shape is constructed from
remains of the original ball, in which the prepupa and later the pupa is formed.
When turning into prepupae the larvae become almost pure white, except for the
mandibles, which are black. Pupal cells were collected during March and beetles
emerged during April.
Newly-emerged beetles are light brown on elytra; the pronotum, head and
legs pale chocolate brown and rather shiny. Up to fifteen pupal cells were
collected from a single cow-pat, usually in association with O. pictus.
Pachnoda sinuata (Fabricius)
DESCRIPTION
Adult (Fig. 20C)
Yellowish beetle with dark-green coloration on pronotum and elytra. Ven-
tral side of body yellow with red and white on sides and about six oval spots on
maroon-coloured apex of abdomen; also two white spots on ventral side of
abdominal apex. Head dark brownish with two long yellowish patches on each
side of clypeus and two oval white spots above eyes. Length about 25 mm. A
variable species of which several varieties have been described.
According to Peringuey (1907) Pachnoda sinuata is widely distributed in
central, eastern and southern Africa. It seems to be fairly well represented in the
western Cape Province, especially in the fruit-producing areas from Worcester to
Somerset West, and has also been collected at Elands Bay. This contradicts
Donaldson’s (1979) statement that it does not occur in the south-western Cape.
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
Soe bina Dia
(i Th,
Z ACN
; ais j) iN Si \ .
gy) HAWERR
ys A/ \ \ MAN
A PG " 4). Mh
Fig. 20. Scarabaeidae. Pachnoda sinuata. A. Pupa, dorsal view. B. Clay cell. C. Adult.
D-I. Larva. D. Apical abdominal segments, ventral view. E. Epipharynx. F. Left maxilla,
dorsal view. G. Mandibles, dorsal view. H. Head, dorsal view. I. Hypopharynx.
SOUTH AFRICAN ARTHROPODS 283
It is fairly common around Milnerton, Pinelands, and even Beaufort West and
Kuruman.
Larva
Very similar to that of Rhinocoeta cornuta (Fabricius) (Fig. 21D) in length,
form and shape, but slightly more hairy on ventral side. Colour of hairs fulvous.
Legs also similar to those of R. cornuta. As in the case of the latter species, they
are able to stretch their bodies fully when walking.
Head (Fig. 20H)
About twice as wide as long, frontal sutures clearly indicated; anterior
extension of coronal suture in most specimens visible as a paler, narrow area.
Ocelli obsolete or faintly visible in some specimens as paler area just behind
antennal bases. Cranium not marbled, or very weakly so; setal pattern almost
same as that of R. cornuta. Antennae about as long as cranium, four-segmented;
apical segment with five oval sensory spots as well as terminal spot with sensory
pegs; penultimate segment also with sensory spot. First and second antennal
segments each with three to four small pits. As in R. cornuta, antennae are bare,
without any setae.
Labrum and clypeus
Labrum nearly twice wider than long; clearly divided into a median and two
lateral lobes. Setal pattern almost the same as in R. cornuta, but with some short
slender setae instead of short spine-like setae on its posterior half. Clypeus about
twice as wide as long and also with same setal pattern as in latter species;
postclypeus also pitted.
Epipharynx (Fig. 20E) very similar to that of Pachnoda impressa (Goldfuss)
(Oberholzer 1959) and R. cornuta, and with about ten strong heli on hapto-
merum; pedium with about twenty strong setae in anterior area, which are almost
as strong as heli; two chaetopariae with fewer setae than in R. cornuta; each
acroparia with about ten strong setae; crepis represented by two slightly sclero-
tized plates. Otherwise as in R. cornuta, except anterior epitorma not so clearly
visible as in latter species and two pairs of macrosensilla transversely arranged in
most specimens examined.
Mandibles (Fig. 20G)
Very slightly longer than cranium and about 1,25 times longer than wide.
Rather similar to those of R. cornuta, but more robust, apical tooth shorter,
particularly on right mandible. Dorsal carina rounded, scrobis obsolete and
represented by two pits, as in latter species, each with seta, distal one longest;
lateral face with about seven setae; brustia as in R. cornuta. Stridulatory area
present ventrally on both mandibles. Cutting edges of both mandibles similar to
those of latter species, but second tooth (ti) on left mandible more acute. As in
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
R. cornuta, a depression present laterad of molar area on each mandible, bearing
about eight fine dorsomolar setae. Otherwise exactly as in R. cornuta.
Maxillae (Fig. 20F)
Very similar to those of R. cornuta. Lacinia and galea fused, demarcating
suture visible dorsally but totally absent on ventral side and separated by strong
sclerotization, part of which is also visible on lateral margin. Galea as in R. cor-
nuta; single uncus present. Dorsally with about six strong setae and about seven
weaker ones exterior to these. On ventral side distally with eight or nine setae,
proximally with only two to three.
Lacinia with two unci, dorsal one with strong spine (yy) at base, which is
more obvious than in R. cornuta. Mesal margin of mala beset with numerous
strong setae or spine-like setae, those bordering unci much more strongly de-
veloped. Stipes ventrally with single long setae near base and distally with about
- three setae on palpifer; latter not clearly demarcated. Dorsally with some fine
setae (about twenty-eight or more) and row of six stridulatory teeth of which
distal one is almost rounded, others acute. Otherwise as in R. cornuta. Maxillary
palp with three distinct segments and basal or fourth segment represented only by
narrow sclerotized plate (y). Penultimate segment with single ventral and lateral
seta. Sensory spot not obvious on apical segment.
Labium
Very similar to that of R. cornuta and with the same setal pattern. Post-
mentum, however, more rectangular. Labial palps as in latter species, with
sensory pegs at apex.
Hypopharynx (Fig. 201) very similar to that of R. cornuta, and with almost
similar setal pattern. However, lateral sclerites less sclerotized in most specimens
seen and small tubercles or sensory pegs form transverse row in front of
hypopharyngeal sclerite; in R. cornuta these are more scattered, forming some-
what curved row.
Ninth and tenth abdominal sterna (Fig. 20D)
The two segments fused with no demarcation line between them on ventral
side. Raster as in R. cornuta, except that there are more palus-like setae on each
tegillum and palidia are almost parallel, each one containing twenty-four to
twenty-six pali. Septum thus formed is long and narrow and usually wider in
middle. Anus and anal lips similar to R. cornuta.
Pupa (Fig. 20A)
Also similar to that of R. cornuta; yellowish brown, but becoming darker as
imago develops. Length 23-25 mm. Apart from differences in developing genital
capsule, pupa of Pachnoda sinuata is easily distinguished from that of R. cornuta
by absence of prothoracic horn and by sinuate hind margin of pronotum (almost
straight in R. cornuta). Each posterolateral angle of pronotum bears large
SOUTH AFRICAN ARTHROPODS 285
rounded tubercle or knob (absent in R. cornuta, and represented only by two oval
folds).
BIOLOGY
In the western Cape Province larvae of Pachnoda sinuata are commonly
found in compost heaps, particuarly in the Milnerton, Pinelands and Somerset
West areas. Large numbers of fairly small and fully mature larvae and pupae were
found during the early winter (April, May) and beetles started to emerge from
about midwinter. However, during the surveys large numbers of larvae remained
in the soil in clay cells until the early spring when pupae were formed, the beetles
emerging from September to November. These surveys indicated that beetles
appear from late autumn to midwinter in the eastern and western Cape as well as
in Natal; the peak period of emergence in the western Cape Province, however,
occurred from early spring to late summer.
The larvae feed on organic material in compost heaps or other suitable
accumulations of decaying organic matter and when fully mature they construct
hard, oval clay cells, 25-30 mm long (Fig. 20B), in which the prepupal and later
the pupal stages are formed. In some instances larvae collected remained for
nearly a month in the prepupal stage before pupation occurred. Pupal stages
lasted for 30—44 days in the laboratory at an average temperature of about 22 °C.
The newly-emerged beetle is pale yellow, marbled with dark green, the head
and pronotum reddish; the ventral side is also pale yellow and reddish. However,
after a few hours to about a day they assume their normal coloration, which is
somewhat darker.
The rearing of Pachnoda sinuata larvae in the laboratory is described by
Donaldson (1979).
Rhinocoeta cornuta (Fabricius)
DESCRIPTION
Adult (Fig. 21A)
Black, pitted all over (particularly pronotum) with setigerous punctures;
areas between punctures shiny, except for the longitudinal matt strips on elytra.
A tubercle or triangular tooth present medially on anterior margin of pronotum
and behind this tubercle usually a wide depression. Legs, ventral side of thorax
and head covered with long golden or fulvous hairs. Length 17-20 mm. For
further details see Peringuey (1907).
Widely distributed in southern Africa (Schein 1960); abundantly found along
the south coast of the Cape Province during present surveys.
Larva (Fig. 21D)
Mature larvae measure about 45 mm in length. Colour is whitish pink to pale
pink, with reddish-brown to light-reddish head. Legs yellow; prothoracic shield
yellowish brown; latter not angled or toothed in front. All spiracles cribriform,
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
/ Ly
Sy ON Wy
Et lise:
Wet :
yy \
Vue 5
Fig. 21. Scarabaeidae. Rhinocoeta cornuta. A. Adult. B. Epipharynx of larva. C. Pupa,
ventral view. D-J. Larva. D. Left lateral view. E. Head, dorsal view. F. Left maxilla,
dorsal view. G. Hypopharynx. H. Apical abdominal segments, ventral view. I. Mandibles,
dorsal view. J. Right mesothoracic leg.
SOUTH AFRICAN ARTHROPODS 287
with concavities pointing as illustrated; thoracic one largest. First seven abdomi-
nal segments with three annulets, eighth with two. Each annulet with broad band
of long spine-like setae. Ninth and tenth terga almost completely covered with
spine-like setae; also some long hairs present. Sterna of abdominal segments 1-8
with row of long hairs and also some shorter spine-like setae; colour of body hairs
fulvous. Legs well developed and more or less of same size; claw of tarsungulus
(Fig. 21J) rounded, fleshy, distally obtuse and surrounded by about ten strong
setae.
The larvae are able to stretch their bodies fully when walking and when
disturbed they usually turn on their backs and slide away from danger.
Head (Fig. 21E)
Slightly wider than long; brownish, smooth; epicranium almost marbled.
Frontal sutures fairly distinct; coronal suture extended anteriorly as paler area,
which sometimes has short branches lying next to frontal sutures. Ocelli present
and clearly visible. Frons with one long seta on each side in most specimens seen,
and two short spine-like paramedian setae just behind frontoclypeal suture, which
is well demarcated; in older specimens with some tiny scattered pits. Also long
seta present on each side near middle of frontal suture. Epicranium with two long
setae dorsally near each ocellus; dorso-epicranial setae represented on each side
by one long and two or more spine-like setae forming longitudinal row surroun-
ded by a narrow paler area, which is not marbled; also some scattered pits, each
with short spine-like seta, which are absent in young specimens. Antennae almost
as long as cranium, four-segmented and in most specimens seen without any
setae. Apical segment with one small round and five large oval sensory spots as
well as terminal sensory spot with sensory pegs; penultimate segment also with
sensory spot.
Labrum and clypeus
Labrum slightly more than half the length of cranium; divided by two clithra
into one median and two lateral lobes. Median lobe with four dorsal setae and
each lateral lobe with about four strong anterior setae and a long posterior one.
Also two long paramedian setae, and in addition some short spine-like setae on
posterior part of labrum. Clypeus about same length as labrum, with one long and
one short seta on each side, as well as two long paramedian setae near posterior
border. Postclypeus abundantly pitted.
Epipharynx (Fig. 21B). Corypha with four strong setae; haptomerum de-
veloped and strongly sclerotized in old specimens. Ten to twelve strong heli
present, situated in a curved row and extended on each side as a curved row of
strong setae that encircles pedium. Latter large, oval, its central area with strong
setae, which are palus-like in front and more weakly developed posteriorly.
Chaetoparia well developed on each side; each acroparia represented by fourteen
to sixteen strong setae; acanthoparia with twelve to fourteen sickle-shaped
setae, last two on each side very small and spine-like. Tormae asymmetrically
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
developed, dexiotorma being largest. Epitormae almost obsolete. Crepis very
weak, area enclosed (haptolachus) with conical sensory cone as well as elongate
plate, which is part of posterior epitorma; fused tormae and anterior epitorma
evident in some specimens examined. Two pairs of macrosensilla longitudinally
arranged. Mesophoba forms continuation of setae encircling pedium and con-
sisting of ten to fourteen fine setae on left side; on right side only about eight setae
present of which inner four to five are stronger developed and spine-like.
Mandibles (Fig. 211)
Mandibles long and narrow (length almost twice width) and somewhat longer
than cranium. Dorsal carina rounded; scrobis weakly developed and almost
obsolete in some specimens, only indicated by two depressions, each bearing
short seta. Six to eight setae also present on each lateral face. About three
small setae (xy) present laterad of molar area, those on left mandible in small
depression. Stridulatory area present ventrally on both mandibles.
Cutting edges of both mandibles fairly narrow and pointed, that of left
mandible with second tooth (ti) rounded, scissorial notch not clearly demarcated
in old specimens; third tooth (tz) more acute, separated by deep notch from
second. Fourth tooth (ts) broad, situated in neck of mandible between cutting and
molar areas. Second tooth (t3) on scissorial area of right mandible fairly broad;
scissorial notch obsolete in old specimens; third tooth (ts) in neck, small and fairly
rounded in some young specimens, more acute in older ones.
Molar area of left mandible clearly divided into proximal (mz) and distal area
(m1); latter broad and forming round lobe-like tooth on ventral side; proximal
area semi-circular posteriorly, with large brustia. Molar area of right mandible
appears trilobed with deep notch between first (ms) and second (ms) and also
between second and posterior tooth (ms). Hind tooth broad posteriorly and
divided by wide concavity into a large dorsal and smaller, more acute ventral
tooth.
Maxillae (Fig. 21F)
Galea and lacinia fused but clearly divided by suture on dorsal side. Galea
with single strong uncus, dorsal surface beset with about twenty-two setae, most
of which are strong and spine-like, particularly those near uncus. Ventral side
with strong spine just below uncus on mesal side and also about seven strong
setae. Lacinia with two unci of which ventral one is smallest; also small spine (yy)
at base of dorsal uncus. Mesal margin of mala beset with numerous strong setae,
of which two are in form of stout spines, one on each side of the two unci. Stipes
dorsally with medium-sized strong setae, particularly on lateral margin, and with
about twelve stridulatory teeth. Ventrally with about eight strong setae distally
and proximally only single strong seta present. Palpifer without any setae. Maxil-
lary palp four-segmented; first segment small and only visible on exterior side;
penultimate segment with one ventral and one lateral seta; apical segment coni-
SOUTH AFRICAN ARTHROPODS 289
cal, with sensory pegs at extreme apex and sensory spot laterally. Labacoria
sclerotized and beset with fine setae on both sides; alacardo with six to eight
setae.
Labium
Distal sclerite of prementum not clearly divided into two lobes and with two
paramedian setae. Proximal sclerite of prementum slightly convex, with two
paramedian setae. Postmentum trapezoidal, with single short seta near each
posterior corner; posterior half pitted in old specimens.
Hypopharynx (Fig. 21G). Glossa beset on each lateral third with numerous
setae, those near anterior lateral corners continuing around palpal bases and
almost encircling labial palpi. In some specimens, sides of median third of glossa
sclerotized; area in between with three strong spines anteriorly; posteriorly on
each side also with about five strong spines and some finer setae, as well as two
median strong spines and about ten tubercles or sensory pegs. Transverse sclerite
strongly developed, asymmetrical and with strong conical tooth on the right as
well as weakly sclerotized area with three or four setae. Lateral sclerites strongly
developed; left one with setae on its lateral and inner margin, right one with setae
only on lateral margin.
Ninth and tenth abdominal sterna (Fig. 21H)
Boundary line between the two sterna hardly visible. Both sterna completely
covered with medium-long, spine-like setae present both ventrally and laterally.
Raster: campus occupied by tegillum; teges separated by two palidia, each consist-
ing of eighteen to twenty-two short pali; septum thus formed wider posteriorly;
same short palus-like setae flank each palidium. Anal opening transverse; lower
anal lip small, not divided. Transverse row of short setae present on either side of
anal slit, which is not visible when viewed from the ventral side.
Pupa (Fig. 21C)
Castaneous brown, first five abdominal segments yellow. However, a few
days before imago emerges, it turns to a dark, almost black colour. Length
21-25 mm. Body almost devoid of setae. Abdominal terga 1—5 with a swelling on
each side. Characteristic conical, median swelling present behind head on front
margin of pronotum, clearly visible in figure.
BIOLOGY
The eggs of Rhinocoeta cornuta are laid during the summer and early autumn
and the larval stage lasts throughout the winter. Mature larvae were already
present in midwinter and those observed during the survey hibernated in the soil,
pupating in the early summer. The larval stage therefore lasts at least 10 months
or more. Observations made along the south coast showed that the larva at first
feeds in the cow-pats and, as it grows bigger, constructs a tunnel in the soil below
290 ANNALS OF THE SOUTH AFRICAN MUSEUM
the pats into which it retreats, only coming up to feed. At first it is bluish white in
colour, but when fully grown it turns pinkish or slightly crimson and is then
40-45 mm long. Larvae collected in June constructed clay cells of about 25 mm
diameter in October, in which they transformed to the prepupal stage. This stage
is characterized by its dirty-white colour and wrinkled appearance, and lasts for at
least 21 days, pupae appearing from about November to January and beetles
emerging from early December to almost the middle of February. The pupal
stage lasts for 25-27 days.
The newly-emerged beetle is light brown on the thorax, with yellow elytra
and abdomen. After about 2 days the thorax becomes black and the elytra reddish
brown and after about 3 days the beetle assumes its normal colour.
Temnorrhynchus retusus (Fabricius)
DESCRIPTION
- Adult (Fig. 14E)
Light brownish red to dark brown in colour, shiny, to slightly shiny; elytra
almost smooth, pronotum fringed with golden hairs, and with fairly large, shallow
pits divided over middle of disc by a narrow glabrous band in most specimens
seen; often with anterior median area of pronotum hollowed out or depressed.
This beetle is easily recognized by the oval flat rugulose or reticulate-rugulose disc
on the head, the edges of which are almost trenchant in some specimens. Length
varying from 14 to 21 mm.
Widely distributed in South Africa and South West Africa (Endrédi 1960).
Larva (Fig. 22A)
Yellowish white in colour. Larva of Temnorrhynchus retusus rather similar to
that of T. coronatus as described and figured by Oberholzer (1963). Thoracic and
eighth to tenth abdominal terga with long slender setae, those of abdominal
segments 1—7 also with short spine-like setae. All abdominal spiracles of about
same size, that of prothorax slightly larger. Legs increasing in length from front to
back, well developed; tibiae of the first two pairs somewhat tapering towards their
apices, those of the hind pairs convex below, not straight (Fig. 22D). Claws of
first two pairs (Fig. 22C) long, somewhat compressed or flattened dorsoventrally
over their distal two-thirds and each with single broad posterior seta near base,
and longer more slender one anteriorly some distance from base, at beginning of
compressed portion. Claws of hind pair (Fig. 22D) short, only about half the
length of those of second pair and each with somewhat sinuate lateral margins
(when viewed from above or below), with single broad posterior seta near base
and somewhat longer seta some distance from apex, as shown in Figure 22E.
According to Crowson (1967) the terga of the ninth and tenth abdominal
segments of the subfamily Dynastinae are completely fused as in the case of the
Cetoniinae. However, in this species the terga are fairly distinct, which agrees
with the drawing by Oberholzer of the T. coronatus larva.
SOUTH AFRICAN ARTHROPODS 291
: oe | at DBE balay 4
ee sal
\
te--< fi == 7p He
Fig. 22. Scarabaeidae. Temnorrhynchus retusus larva. A. Left lateral view. B. Head, dorsal
view. C. Tibia and tarsus of mesothoracic leg, posterior view. D. Tibia and tarsus of meta-
thoracic leg, posterior view. E. Claw of metathoracic leg, seen from below. F. Apical abdomi-
nal segments, ventral view. G. Anterior margin of labrum, viewed directly from the front.
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
Head (Fig. 22B)
Slightly less than one-third wider than long, sides and hind margin very
convex; reddish to brownish in colour, almost superficially rugoso-reticulate,
forming three inconspicuous ridges on frons, separated by areas of lighter sclero-
tization; first transverse area just behind frontoclypeal suture fairly deeply exca-
vated in some specimens and bearing about six anterior frontal setae. Second
light-coloured area bears two exterior frontal setae on each side as well as two
paramedian setae. In this species four posterior frontal setae present behind third
transverse ridge; otherwise chaetotaxy as illustrated. Frontal and coronal sutures
faintly visible.
Four-segmented antennae about half as long as width of head; first, third and
apical segments of about equal length, second segment longest. Apical segment
with large dorsal oval sensory spot as well as two smaller spots on ventral side;
penultimate segment also with oval sensory spot on cone-shaped apical projec-
tion.
Labrum and clypeus
Labrum pitted, about one-third wider than long, narrower and about one-
third longer than clypeus; its anterior and lateral margins convex. Anterior
transverse row of four setae present, one of which is situated on each lateral
margin; posterior row of about eight setae as illustrated. Also two spine-like setae
present in middle of anterior margin. Clypeus nearly two-thirds wider than long,
its anterior margin slightly concave. Postclypeus also pitted with transverse row of
four setae, one of which is situated on each lateral margin; also a shorter lateral
seta near anterior margin of postclypeus.
Epipharynx (Figs 22G, 23A). Corypha fairly well indicated, with about five
strong setae on its ventral margin and about seven bordering the dorsal edge.
Epizygum fairly discernible in most specimens seen. Haptomerum strongly
sclerotized and with about seven sensilla and three broad teeth or heli flanked on
right side by about eight strong dentiform setae of the chaetoparia. Latter beset
with strong setae on both sides, the setae diminishing in size towards the plegma-
tia, forming pitted gymnopariae. Acropariae each with about fifteen long setae of
which four to five are situated in a dorsal position. Each acanthoparia with ten to
twelve sickle-shaped setae; each seta with indistinct plegma. Pedium well indi-
cated, traversed in middle by narrow anterior epitorma. Both dexio- and
laeotorma as narrow strongly sclerotized plates; laeotorma with small almost
conical pternotorma, which is rather similar to that of Rhinocoeta cornuta.
Mesophoba on each side as row of slender setae. Crepis indicated by fold,
haptolachus includes an almost triangular sensory plate, a sensory cone and two
macrosensilla on each side.
Mandibles (Fig. 23E-—F)
Fairly long, about one-third longer than wide. Dorsal carinae rounded and
scrobis with one medium-long, broad distal seta and longitudinal row of five small
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294 ANNALS OF THE SOUTH AFRICAN MUSEUM
pits, each with short seta. Lateral areas with ten to fourteen setae; also about
seven setae at base on ventral carina. Medium-long seta present in depression on
neck of scissorial area, about half-way between seta on scrobis and third tooth
(t2 and ts).
Cutting edge of left mandible with first and second tooth (t:) united as broad
blade, with deep notch between second and third tooth (tz). Small almost truncate
tooth (ts) present at junction of molar and cutting edge on left mandible, absent in
right one but represented by distinct tooth (ts), present about half-way between
rounded third tooth (ts) and molar area. Second (ts) and third tooth (ts) on right
mandible rounded, in most specimens rather indistinct, in some almost obsolete.
Molar area on right mandible smaller than that on left, with upper strong
tooth (ms) forming transverse ridge in front of which is fairly distinct ventral tooth
(mc). Second molar tooth (ms) small, also forming ridge, similar to that of m3 and
strongly developed on ventral side. Posterior tooth (ms) almost as broad lobe,
- bearing brustia. Molar area of left mandible with broad lobe-like transverse tooth
(m1) followed by second conical tooth (m7). Proximal molar area (mz) in this case
broad, rounded posteriorly with almost semi-circular row of about twenty-two
strong setae, and forming a short broad anteroventral tooth as well as a broad,
conical anterodorsal tooth, behind which is almost triangular acia with four short
apical setae. Also row of about seven dorsomolar setae present on inner margin
of both molar areas.
Ventrally each mandible with broad, oval stridulatory area with small pit
carrying two setae, about half-way between it and molar area (this pit with setae
also present in most other scarab larvae).
Maxillae (Fig. 23H, J)
Galea and lacinia united, but clearly indicated by dividing suture on dorsal
side, ventrally only by S-shaped sclerotized line. Galea with single long acute
uncus at apex, dorsally with three strong setae below uncus and longitudinal row
of three to four more slender setae along dividing suture, as well as some smaller
slender setae; ventrally also with three strong setae below uncus as well as some
more slender setae as indicated in figure; strong palus-like seta present on base of
uncus on lateral margin. Lacinia with three conical unci, somewhat smaller than
that of galea, bordered by strong broad setae, its mesal margin fairly densely
covered with strong setae. Stipes dorsally with about six truncate stridulatory
teeth and distad of these obliquely placed rounded and wider stridulatory
tubercle; ventrally and dorsally with fine slender setae, some also on lateral
margin and single proximal one close to border of disticardo.. Palpifer clearly
demarcated, without any setae and bearing four-segmented palp of which first
segment is shortest; third segment somewhat longer than second and with single
ventral and lateral seta; fourth segment longest, about as long as second and third
together, and with some sensory cones at apex. Both disti- and proxicardo with
some setae, ventrocardo without setae. Labacoria fairly convex ventrally and
beset with slender setae on both sides.
SOUTH AFRICAN ARTHROPODS 295
Labium (Fig. 231)
Postmentum slightly wider than long, with single short seta near each pos-
terior corner; proximal sclerite of prementum wider than long, convex, with
slender setae; distal sclerite with two long paramedian setae near palpal bases,
and some slender setae on each side. Two-segmented palpi with apical segment
almost twice as long as first and with sensory pegs at apex.
Hypopharynx (Fig. 23G). Glossa convex, its central disc with short spine-
like setae, arranged almost in a circle in some specimens; on each side with
slender setae. Hypopharyngeal sclerite strongly sclerotized, asymmetrical, with
strong tooth with small depression bearing brush of fine setae on right side; more
weakly sclerotized on left. Lateral sclerites less sclerotized, lateral margins acute
or pointed in middle, on left with some setae distally, also curved row of fine setae
down its middle and along posterior lateral margin; on right side also with some
fine setae distally near base of transverse tooth and row near basal border of
latter.
Ninth and tenth abdominal sterna (Fig. 22F)
Dividing line between two segments clearly indicated. Ninth sternum with
transverse row of four setae near posterior border, two of which are more or less
paramedian setae, situated fairly close together. Tenth sternum also with four
setae, the median ones far apart. Raster: teges with about thirty-five to forty
hamate setae; palidium absent; also thirty-five to forty hamate setae on lower anal
lip and some on upper anal lip. Internal sclerotized pouch on venter of
ninth abdominal sternum similar to that described by Oberholzer (1963) for
T. coronatus.
Pupa (Fig. 23B—-D)
Castaneous brown and rather similar in colour to that of Rhinocoeta cornuta.
Easily recognized by flat disc on the head as in adult. Hind margin of pronotum
almost straight as in R. cornuta. Spiracles also similar to those of both the latter
and Pachnoda sinuata, first four pairs large and oval, first pair covered by wing
sheaths; fifth to eighth pair smaller and inconspicuous. Pupa of this species lacks
the five pairs of dorsal abdominal tubercles or swellings present in the case of
R. cornuta and P. sinuata. Legs with tarsal sheaths much shorter than in other
two species. Developing genital capsule similar to that of P. sinuata, but three
swellings much more prominent than in latter. Length 23-25 mm.
BIOLOGY
Larvae of Temnorrhynchus retusus were found in sandy soils in the Cape
Peninsula throughout the year, feeding on the roots of grasses and other garden
plants. Pupae formed in clay cells were observed during November and the
beetles emerged during December. Newly-emerged beetles are brownish-red to
reddish, but assume their normal coloration within a day or two.
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Trox rhyparoides Harold
DESCRIPTION
Adult (Fig. 24A)
Peringuey (1901) and Scholtz (1980) give a good description of Trox rhypar-
oides. Blackish, but usually covered with soil which gives it a greyish appearance.
Head and thorax closely pitted. Clypeus triangular and pointed in front.
Prothorax raised in middle; this raised area wider in front than behind, with three
impressions in middle and wider one behind; disc on each side behind this raised
area with median carina. Head also with strong transverse ridge intersected in
middle by blunt carina which ends at apex of pointed clypeus. Elytra with about
eight strong costae, area between them with two rows of foveae with tiny, shiny
tubercles in between. Head and prothorax fringed with spoon-shaped hairs.
According to Scholtz (1980) the absence of setal tufts and fringes on the
_pronotum and elytra and the distinct costal ridges make 7. rhyparoides a very
distinctive species. In specimens examined the lateral margins of elytra have only
very small spine-like setae and the fosciculate tubercles, present in other species
of Trox collected during the surveys, are absent. Length 7,1-8,5 mm.
Widely distributed in the Republic of South Africa and Lesotho, and also
present on St. Helena (Scholtz 1980).
Larva (Fig. 24D)
Baker (1968) gives a general description of the larvae of the genus Trox. In
T. rhyparoides mature larvae are almost white with golden-brown head, which is
somewhat piceous on the sides above eyes. In young stages, head 1s pale piceous
red. Abdominal segments 1-8 divided into three annulets, those of segments 1—6
and first two annulets of segment 7 bearing short, spine-like setae as well as long
hairs. Eighth segment devoid of spines. Prothoracic shield fairly clearly indicated,
its front margin straight. Spiracles (Fig. 24M) biforous; under high magnification
three sclerotized struts clearly visible, with middle one somewhat stronger than
other two. According to Hinton (1967) the trogids have elateroid spiracles which
evolved from the primitive type of spiracle in which the orifice of the spiracle is
not sufficiently blocked by cuticular struts to prevent the withdrawal through it at
ecdysis of the old spiracle and tracheae. Legs well developed and covered with
long hairs; claws (Fig. 24E) acute, simple, each bearing two medium-long setae
near base.
Small larvae measuring 5,5 mm in length are very similar to full-grown
specimens, but differ from the latter in the smaller number of annulets which have
spines; in specimens examined only one annulet on each of abdominal segments
1-7 contained spines.
Head (Fig. 24G)
Wider than long; frontoclypeal suture obsolete in middle and forming wide
U-shaped depression with four hairless pits. Frons deeply pitted; four of pits on
SOUTH AFRICAN ARTHROPODS 297
LEE SWIPE
Fig. 24. Scarabaeidae. Trox rhyparoides. A. Adult. B. Pupa, ventral view. C-—E. Larva.
C. Apical abdominal segments, ventral view. D. Larva, left lateral view. E. Right mesothoracic
tibia. F. Apex of pupa, left lateral view. G—M. Larva. G. Head, dorsal view. H. Epipharynx.
I. Hypopharynx. J. Left mandible, dorsal view. K. Right mandible, dorsal view. L. Left
maxilla, dorsal view. M. Biforous abdominal spiracle (highly magnified).
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
each side with long seta each. Each side of epicranium with four dorsal setae, of
which dorso-epicranial one is often double; some pits also hairless. Ocelli present
and fairly prominent behind antennal bases. Antennae much shorter than
cranium, three-segmented, third segment very small.
In young specimens, measuring 3—3,5 mm, posterior epicranial (exterior
dorso-epicranial) setae are absent; otherwise head is similar to that of mature
specimens.
Labrum and clypeus
Labrum about as long as clypeus. Its anterior margin not clearly divided by
weakly developed clithra; dorsally with six long setae along front margin, as well
as two paramedian setae on central disc. Postclypeus with only two setae.
Epipharynx (Fig. 24H). Plegmata and proplegmata absent. Corypha with
four strong setae. Acanthoparia with five to six almost sickle-shaped setae.
_Acroparia represented by three asymmetrically arranged setae on each side of
haptomeral area; two long setae (close together) present on right side near
anterior border; on left, however, one long seta situated near pedium and one
near anterior border; small seta posteriorly on each side of haptomeral area;
these setae may be longer in older specimens. Haptomerum almost conical, fairly
well sclerotized in old specimens and without teeth in specimens examined,
except for two small setae proximally and two sensilla. Pedium raised and convex,
central area with stout hairs or groups of spines, particularly on proximal area;
distally with short spines and curved row of twelve to fourteen sensilla anterior to
these. Tormae (dt and It) symmetrically developed with short anterior epitorma.
Chaetoparia represented by two setae proximally along anterior arms of tormae.
Crepis represented by two small sclerotized areas; sclerotization more complete
in older specimens; two sclerotized or sensory plates and sensory cone also
present.
Mandibles (Fig. 24J-K)
Slightly more than half the length of cranium, both mandibles slightly longer
than wide. Dark golden brown in colour with molar, scissorial and lateral edges
black. Neck of both mandibles short; scrobis and dorsal carina well developed;
scrobis with one long proximal and shorter distal seta. Stridulatory area absent on
both mandibles, but each with small brustia.
Left mandible with broad blade-like cutting edge; scissorial notch indicated
in most specimens; third tooth (t2) small and acute in some specimens. Cutting
edge of right mandible also fairly broad, with two distinct teeth; second tooth (ts)
broad, scissorial notch well marked in most specimens.
Molar areas of both mandibles divided into two teeth, of which distal tooth
(m; and m3) is conical and situated almost half-way between last scissorial tooth
(t2 or ts) and proximal molar area (m2 or ma). Tiny longitudinal groove with fine
dorsomolar setae present on both mandibles.
SOUTH AFRICAN ARTHROPODS 299
Maxillae (Fig. 24L)
Galea and lacinia free for most of their length. Galea bearing single strong
uncus; lacinia with two strong unci and third smaller tooth on ventral side. Galea
with five strong setae around uncus of which two are situated ventrally. Lacinia
with about thirteen strong setae on mesal margin, arranged more or less in two
rows. Stipes with only two long setae, one on ventral side near cardo and one on
palpifer; dorsally without setae, but with strong ridge (c) bearing about twenty
stridulatory teeth. Cardo with only one short seta on disticardo; labacoria beset
with minute spinules. Maxillary palp four-segmented, with two setae on ventral
side of penultimate segment and one on ventral side of basal segment. Palpifer
not clearly demarcated on dorsal side and bearing few small stridulatory teeth.
Labium
Submentum trapezoidal, with single seta on each side near lateral margin.
Mentum convex, with two long setae. Distal sclerite of prementum not divided
into lobes but with two setae near anterior margin, one on each side of palpiger.
Two setae also present posteriorly near border of proximal sclerite of premen-
tum. Palpiger also with some small spine-like setae on external border.
Hypopharynx (Fig. 241). Distal region of glossa with only four setae and
some minute spines or cones, which are in some specimens almost semi-circularly
arranged; also with semi-circular row of fine hairs on each side. Hypopharyngeal
sclerite weakly developed and with row of fine hairs in front and on each side; in
older specimens hypopharyngeal sclerite more strongly developed, particularly its
two anterior arms (subapotormae). Same applies to lateral sclerite (subtormae).
Ninth and tenth abdominal sterna (Fig. 24C)
Dividing line between these segments lacking: ninth sternum with transverse
row of about eight setae, tenth with transverse row of about twelve. Raster
absent. Upper anal lip small, lower anal lip almost semi-circular and divided by
slit into two halves. The feebly trilobed anal area suggests relationship to the
stag-beetles, Lucanidae (Hayes 1929).
Pupa (Fig. 24B)
Newly formed pupa yellowish white and 8—9 mm long. Anterior and lateral
borders of pronotum as well as anterior part of head setose. Meso- and
metathorax dorsomedially bearing two closely approximated tubercles, each one
with about four setae. Six fleshy protuberances also present laterally on abdomen,
each bearing five to seven long setae. Apex of abdomen with long setae and two
fairly long hooks pointing upwards (Fig. 24F).
BIOLOGY
Trox rhyparoides beetles were found in large numbers under decaying ani-
mals around Cape Town from March to November. They feed on bits and pieces
of flesh left on the skeletons as well as on loose hairs scattered on the ground
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
around the carcasses. Young larvae occurred during May and again during
October and pupae were collected around the middle of August and also at the
end of October. According to available information, the larval period lasts for
about 6 weeks and the pupal stage 10-14 days.
ACKNOWLEDGEMENTS
I am deeply grateful to Dr V. B. Whitehead and Miss M. Macpherson, both
of the South African Museum, Cape Town, for their comments. Also special
thanks to Mr V. Branco of the same institute for preparing some of the figures.
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LAING, F. 1928. A note on two parasites of Lyctus parallelopipedus Mel. The Entomologist 61:
163-164.
Le Pettey, R. H. & Gopparpb, W. H. 1952. On the control by insecticides of Heteronychus
consimilis Kolbe (Dynastinae), a serious pest of wheat in Kenya. Bull. ent. Res. 43: 402—406.
Morison, G. D. 1925. The Khapra beetle (Trogoderma granarium Everts). Proc. R. phys. Soc.
Edinb. 21: 10-13.
OBERHOLZER, J. J. 1958. A description of the third stage larva of Onitis caffer Boh. (Copridae:
Coleoptera) with notes on its biology. S. Afr. J. agric. Sci. 1: 415-422.
OBERHOLZER, J. J. 1959. A morphological study of some South African lamellicorn larvae. 1.
Description of the third instar larvae. S. Afr. J. agric. Sci. 2: 41-74.
OBERHOLZER, J. J. 1963. Description of the larva of Temnorrhynchus coronatus F. (Dynastidae:
Coleoptera). S. Afr. J. agric. Sci. 6: 85-90.
PERINGUEY, L. 1901. Descriptive catalogue of the Coleoptera of South Africa. Trans. S. Afr.
phil. Soc. 12: 1-560.
PERINGUEY, L. 1907. Descriptive catalogue of the Coleoptera of South Africa. Trans. S. Afr.
phil. Soc. 13: 294-546.
PETERSON, A. 1967. Larvae of insects. Part 2. Coleoptera, Diptera, Neuroptera, Siphonaptera,
Mecoptera, Trichoptera. Columbus, Ohio: Edwards Brothers Inc. (Lithographed. )
Prins, A. J. 1965. Notes on the biology and morphology of the wattle chafers Monochelus
calcaratus Burm. (Melolonthidae), Hypopholis sommeri Burm. (Melolonthidae), and
Adoretus ictericus Burm. (Rutelidae) with some references to natural enemies (Coleoptera:
Lamellicornia). Entomology Mem. Dep. agric. tech. Servs Repub. S. Afr. 9: 1-55.
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
REES, B. E. 1943. Classification of the Dermestidae (larder, hide and carpet beetles) based on
larval characters, with a key to the North American genera. Misc. Publs U.S. Dep. Agric.
511: 1-18.
Rose, D. J. W. 1964. A new surface beetle Herpiscius somneri Sol., a pest of maize. Rhodesia
agric. J. 61: 29.
SCHEIN, H. 1954. Uber Coenochilus (Coleoptera, Cetonitidae). Eine Revision der afrikanischen
Arten mit einer Bestimmungstabelle und ein Nachtrag zu meiner Revision der asiatischen
Arten. Ent. Arb. Mus. Georg Frey 5: 271-325.
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83-112.
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Africa. Cimbebasia Mem. 4: 1-104.
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appl. Biol. 6: 101-115.
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30: 845-863.
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Trustees, British Museum (Nat. Hist.).
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Voss, E. 1973. Coleoptera Curculionidae partim. S. Afr. anim. Life 15: 395-479.
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WINKLER, J. R. 1960. Coleoptera Cleridae. S. Afr. anim. Life 7: 127-162.
WitIlMER, W. 1960. Coleoptera: Cantharidae, Malachiidae and Dasytidae. S. Afr. anim. Life 7:
113-127.
ABBREVIATIONS
ac alacardo
aca acia
acp acanthoparia
acr acroparia
aet anterior epitorma
an anus
ant antenna
br brustia
c; carc carina; ridge
car cardo
cl clithrum
co corypha
cos epicranial suture; coronal suture
cpa chaetoparia
cpp caudal support projection
cr crepis
csc convex sclerite of cardo
CX coxa
hps; hsc
ial
SOUTH AFRICAN ARTHROPODS
disticardo
dorsal carina
dorso-epicranial setae
distal sclerite of prementum
dexiophoba
dexiotorma
compound eye
exterior anal lobe
exterior dorso-epicranial setae
epicranium
epipharyngeal sclerome
epizygum, zygum
frons
finger-shaped air-tube of biforous spiracle
femur
developing genital capsule
galea
glossa
protrusible gland
gymnoparia
gin trap
gula
head
haptomerum
hypopharyngeal sclerome; hypopharyngeal sclerite
interior anal lobe
opening of internal sclerotized pouch
claw; tarsungulus
juxtastipes
ligula
labacoria
labium
labrum
lacinia
lower anal lip
lateral lobe setae
prostheca
labial palp
laeophoba
labial stipes
lateral sclerite
laeotorma
mentum
molar areas
mala
mandible
superlinguae; maxillulae
median lobe setae
maxillary palp
mesophoba
macrosensillum
maxilla
ocellus
occipital foramen
oncyli
scissorial area
palidium
paramentum
postclypeus
precardo
pedium
303
ANNALS OF THE SOUTH AFRICAN MUSEUM
penicillus
posterior epitorma
palpifer
palpiger
prementum
postmentum
proxicardo
proximal sclerite of prementum
protophoba
pteronotal support projection
pternotorma
retinaculum
spur
subapotorma
sensory cone
subcardo
scrobis
stridulatory teeth
sensory spot
sensory punctures
septum
straining hairs
sensory appendix; supplementary joints of antennae
submentum
scissorial notch
subnasal sclerite
sensory plate
spiracle
maxillary stipes
subcardo
tarsungulus
mandibular teeth; hypopharyngeal teeth
tibia
teges
tegillum
torma
dorsal tergal support projection
transverse ridge
tubercle
upper anal lip
uncus
urogomphus
ventrocardo
short and long setae on labrum of scarab larva
dorsomolar setae
narrow sclerotized plate
spine at base of uncus
tuft of hairs
median hairs near labral apex
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
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punctuation and not be abbreviated; if the year is added, a comma must separate author’s
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Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
Figs 14-15SA
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
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figures of plates are enclosed in parentheses to distinguish them from text-figures
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the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new species, One specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus...’; *. . . in C. namacolus (Fig. 10)...’
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by initials or full names
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Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
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book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
‘Biological Abstracts.
A. J. PRINS
MORPHOLOGICAL AND BIOLOGICAL
NOTES ON SOME SOUTH AFRICAN
ARTHROPODS ASSOCIATED
WITH DECAYING ORGANIC MATTER
PART 3
THE FAMILIES DERMESTIDAE,
CANTHARIDAE, MELYRIDAE,
TENEBRIONIDAE, AND SCARABAEIDAE
(COLEOPTERA)
—
i...
QH rar ms 94 PART 5 DECEMBER 1984 ISSN 0303-2515
CAPE ‘TOWN
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5. REFERENCES cited in text and synonymies should all be included in the list at the end of
the paper, using the Harvard System (ibid., idem, loc. cit., op. cit. are not acceptable):
(a) Author’s name and year of publication given in text, e.g.:
‘Smith (1969) describes .. .’
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‘As described (Smith 1969a, 1969b; Jones 1971)’
‘As described (Haughton & Broom 1927)...’
‘As described (Haughton ef al. 1927)...’
Note: no comma separating name and year
Dagination indicated by colon, not p.
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within each name, with suffixes a, b, etc. to the year for more than one paper by the same
author in that year, e.g. Smith (1969a, 19695) and not Smith (1969, 1969a).
For books give title in italics, edition, volume number, place of publication, publisher.
For journal article give title of article, title of journal in italics (abbreviated according to the World list o,
scientific periodicals. 4th ed. London: Butterworths, 1963), series in parentheses, volume number, part
number (only if independently paged) in parentheses, pagination (first and last pages of article).
Examples (note capitalization and punctuation)
BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.
FIscHER, P.-H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.
FIscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs
Zool. exp. gén. 74: 627-634.
Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.
Ann. Mag. nat. Hist. (13) 2: 309-320.
Koun, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.
Bull. Bingham oceanogr. Coll. 17 (4): 1-51.
THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische
und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.
Jena: Fischer. Denkschr. med.-naturw. Ges. Jena 16: 269-270. :
(continued inside back cover)
ANNALS OF THE SOUTH AFRICAN MUSEUM
ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
Volume 94 + 2Band
December 1984 Desember
Part 5 Deel
A NEW SEA ANEMONE FROM SOUTH AFRICA
(ANTHOZOA, PTYCHODACTIARIA)
By
K. W. ENGLAND
&
E. A. ROBSON
Cape Town Kaapstad
The ANNALS OF THE SOUTH AFRICAN MUSEUM
are issued in parts at irregular intervals as material
becomes available
Obtainable from the South African Museum, P.O. Box 61, Cape Town 8000
Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM
word uitgegee in dele op ongereelde tye na gelang van die
beskikbaarheid van stof
Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad 8000
OUT OF PRINT/UIT DRUK
i GES) SO) I, OS, ee SE), 5, ED).
GOs tes TOD), ©, CUD, M), OS).
11(122,5, 7et-=pac), 14-2), 15425), 24(2),127, 3110-3), 32), 33, 26) 45)
Copyright enquiries to the South African Museum
Kopieregnavrae aan die Suid-Afrikaanse Museum
ISBN 0 86813 062 1
In Suid-Afrika gedruk deur
Die Rustica-pers, Edms., Bpk.,
Courtweg, Wynberg, Kaap
Printed in South Africa by
The Rustica Press, Pty., Ltd.,
Court Road, Wynberg, Cape
A NEW SEA ANEMONE FROM SOUTH AFRICA
(ANTHOZOA, PTYCHODACTIARIA)
By
K. W. ENGLAND
&
E. A. ROBSON
Department of Pure & Applied Zoology, University of Reading
(With 11 figures and 2 tables)
[MS accepted 23 May 1984]
ABSTRACT
Preactis millardae gen. et sp. nov., found typically off the Atlantic coast of the Cape
Peninsula, is described with details of its distribution and behaviour. The species has taxonomic
characters that compare with those of Dactylanthus antarcticus Carlgren, 1911, and place it in the
order Ptychodactiaria, but it also possesses characters that necessitate a redefinition of that order
and the creation of a new family, the Preactiidae, to accommodate it. Some additional data are
also given on D. antarcticus.
CONTENTS
PAGE
MMUHOCUC HOM Gets Neat se yee are waar IMT on A oe he sae 306
©ccurence andigeneraliobservations) 4.44545 6494024 eee 306
omanvRock. FalseiBaye y. tiaras sy octat toe seme een 306
Onrus near Henmanuse. 9). oe oe ee ee ae 309
@Oudekraals@ape Reninsulayn. = os sane eae ae 309
Maclear’s Beach, Cape Point Nature Reserve .......... 310
Atlanticicoastot the Cape Peninsula =. 224.24). 4-4-52-- 310
Maxonomic descnption (by K7 WeEngland)*..3-3225. 9.45 ee: S15)
Order Piychodacharia (redefined)!--4).245--oe saeco: 315
FamilyeRreactidae famianOVvere 4444. esa ee ae oe SS
eRCACHSIC EM MOV ARE © enh: Geta deel ern Saudi. ha cease, oe 316
LR CACKSHMUNAKAGe Sp aNOVn nee ae eee) Aetna ee oe 316
Wiatenialiptacenters cht cette ce ee ae chen eee 316
Cy MOlO Sy ae terns elite oan edatcras Sere EES 2 SMU7/
DeSCHIpPlOndeses eos ee eer Fo 8 ah Ree aaron - Sly
PRIVAT OMY ea hr ets cca soiree tie SS peat Og S77,
INIESEMLETICS i eae ctare ert nerey tac eee alah aid ee ne: 319
Gaim ee Tae aha tease ee EE ee an hohe ia aoe a B22
IdlemtiticatiOneexcnt ea wae. 2 hates ean ne ey 323
COMMITS MESH ecm cnc ie ated apie s, ulanes 326
PACKNOWICESEMETISE seers itv aida cihea cabelas acta ey 2 chaos Seweocas 328
INGLES LESS sac enue es. vai ccs Cha is chia eneus t eeee Coreen tame 328
INCIETENCE SG. Meek on Uae EPS. PEt See aes eke remeee ee 329
305
Ann. S. Afr. Mus. 94 (5), 1984: 305-329, 11 figs, 2 tables.
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
INTRODUCTION
This paper describes a South African sea anemone that is a most interesting
addition to a rare and relatively little-known group, the Ptychodactiaria. Until
now the Ptychodactiaria have included one species recorded from the Antarctic
and one from the Arctic (Carlgren 1949). The description of this new species is
introduced with a summary of what is known about its occurrence and biology,
together with available photographic records. It is hoped that this will stimulate
further discoveries.
Apart from three finds of specimens washed up on beaches, SCUBA divers
have discovered the new anemone attached to rocks at depths of 10—20 m. It has
been found at Oudekraal off the Atlantic coast of the Cape Peninsula, which
would seem to represent a typical habitat (Velimirov et al. 1977), and at Roman
Rock in False Bay. Stranded specimens have been found at Maclear’s Beach
(near Cape Point) and near Hermanus. This is undoubtedly a cold-water species
‘that occurs in the Benguela Current (see Day 1970; Brown & Jarman 1978;
Andrews & Hutchings 1980; Field et al. 1980). Its origin and precise distribution,
however, remain unknown. It is worth noting that specimens so far discovered all
seem to have been of adult size, and that small anemones or juveniles, which
might be expected to occur in a representative cross-section of the population,
have not yet been found. Of seven preserved individuals all except the smallest
one bear gonads, but nothing is yet known about the life history or about possible
modes of dispersal.
OCCURRENCE AND GENERAL OBSERVATIONS
During the period 1972—82 several specimens were collected and information
about their distribution and general biology is drawn up here in chronological
order. Localities indicated on the map (Fig. 1) are sites corresponding to five sets
of records.
ROMAN ROCK, FALSE BAY
Two specimens at about 10 m on red gorgonians were collected by SCUBA
diving and brought by R. J. Griffiths to J. H. Day, 3 December 1972. Preserved
specimens now mislaid (CP 809, Zoology Department, University of Cape
Town), but identifiable from Day’s notes and two photographs taken by him:
‘Basal disc poorly developed, not attached. Animal mobile, progresses by means
of hollow finger-like appendages on column wall. Oral disc smooth no tentacles
visible—maybe a few small tentacles inside mouth—requires dissection. Thin
translucent body wall—white, column appendages white with yellow-brown tips.
Brown radiating lines on basal and oral discs which appear to mark insertion of
mesenteries. 2 siphonoglyphs only.’
One of the photographs is shown in Figure 2A.
307
A NEW SEA ANEMONE
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W pur [eeryopng ‘sniuO *‘yooy ueWoY 1X9} 94} Ul 0} postojor suoumIoads Jo says UOKATTOO suimoys dey *[ “314
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308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 2. A. Specimen from Roman Rock, False Bay (CP 809, photo J. H. Day). Note mesentery
insertions. Approx. X 1. B-—D. Specimen from beach at Onrus (SAM-—H1677, photos S. X.
Kannemeyer). B. Note peristaltic waves (moving from the base) and vesicles in transverse rows.
Approx. X 1. C. Oral disc, x 2. D. Pedal disc, x 1,5.
A NEW SEA ANEMONE 309
ONRUS, NEAR HERMANUS
One specimen found loose in a rock pool, collected by M. R. Polchet and
brought to N. A. H. Millard at the South African Museum, 4 February 1973. This
anemone was observed and preserved by Millard (SAM-—H1677), and photo-
graphed by S. X. Kannemeyer of the South African Museum. The photographs
suggest that the column was up to 13 cm long, the basal disc about 5 cm and the
oral disc 1,7 cm in diameter. General coloration orangey-pink: the tips of the
vesicles covering the column darker in tone, the column pale. Mouth and oral
disc, also the pedal disc, white with twenty-four radial red stripes. The vesicles
appeared to be in transverse rows but tentacles were not evident. Millard also
noted that the animal made no attempt to attach to the dish by means of the base
or vesicles, and performed ‘peristaltic’ contractions of the whole column. Peristal-
tic waves moved distally from the base.
The photographs are shown in Figure 2B-D.
Millard’s further comments: ‘It was probably brought into this coast by
currents or wind’ (7 May 1973); and ‘I have shown the slide to various people and
am told that this creature has been seen before several times’ (11 April 1975).
OUDEKRAAL, CAPE PENINSULA
A cold-water habitat at about 10-12 °C, very exposed conditions with fast
current, depth 10-15 m. About six specimens seen while SCUBA diving by
C. L. Griffiths and the two largest were collected on 7 October 1976. All
appeared to be ‘attached’ to a vertical granite rock by the pedal disc: they were
vertical in position and ‘closed up’. Griffiths observed some variation in size, from
perhaps 5 cm upwards, although he thought that apparently smaller specimens
might simply have been more contracted. There was some variation in colour
between individuals, but the two collected looked almost identical. The habitat is
characterized by Allopora, gorgonians (Eunicella), solitary corals, golf-ball
sponges (Haliclona) and Gorgonocephalus, as described by Velimirov ef al.
(1977).
The two specimens collected (SAM-—H2822 and BMNH 1983.4.27.1) were
observed and photographed in the Zoology Department, University of Cape
Town, by Griffiths and by Robson, who preserved them after making the follow-
ing notes.
In one specimen the column was at least 20 cm long, the pedal disc 7-9 cm
and the oral disc 2 cm in diameter. Vesicles covering the column pinkish mauve
with orange tips. Twenty-four radial stripes marking the insertion of mesenteries
dark red on the oral disc, orange on the pedal disc. On the column, brownish lines
correspond to mesenteries. Background colour pale orange. On the column,
vesicles are of all sizes and seem to lack regular alignment. There are two
siphonoglyphs, and bilateral symmetry of the pedal disc is also evident.
In the laboratory detached specimens readily change shape. Their mobility of
profile gives the impression that behaviour in natural conditions may be fairly
310 ANNALS OF THE SOUTH AFRICAN MUSEUM
adaptable. Relatively little, however, was discerned as the specimens observed
were subjected to bright light and to higher temperatures than usual. This
anemone exhibits no specialized rapid contractions. Peristaltic waves were seen to
travel quite fast, in this case from the oral disc to the base. The circular muscle of
vesicles as well as of the column may be involved. The specimen in Figure 3A
reacted to a mechanical stimulus with two or three peristaltic waves after which
the column shortened to about 10 cm. When the column was contracted the pedal
disc usually became inflated and very thin but it showed no tendency to adhere.
The other specimen collected by Griffiths, however, soon afterwards attached
itself quite firmly by the base to the bottom of a deep plastic bowl, assuming the
shape of a truncated cone. In both specimens the vesicles throughout seemed
non-adhesive and they did not move individually unless touched. In general their
changes in length or tone were associated with those of other muscles (e.g., as
noted, peristalsis of the column) and local responses of their circular and longitu-
dinal muscles were not very evident. In the Oudekraal habitat some ability to
reattach to rocks if swept off by the current may be expected, but laboratory
observations provided few clues about this.
Griffiths noted in a letter: ‘Incidentally, the anemone is quite common—
I have mentioned it to several divers and they all know about it’ (2 May 1980).
Photographs by Griffiths of one of the specimens are shown in Figure 3A—D.
The photograph for Figure 4A was taken by Robson.
MACLEAR’S BEACH, CAPE POINT NATURE RESERVE
Four small specimens found stranded by S. X. Kannemeyer on 4 August 1980
are now in the South African Museum (SAM-—H3030). Preserved in alcohol,
three of these specimens measure 3, 4 and 5 cm in length respectively.
One stranded specimen found in a tide pool by T. M. Gosliner on
27 November 1981. He noted ‘very bright orange coloration’.
Cnidae from SAM-—H3030 are shown in Figure 4B.
ATLANTIC COAST OF THE CAPE PENINSULA
Observed by T. M. Gosliner to occur commonly at 10-20 m, e.g. at Hotten-
tot’s Huisie, near Oudekraal, during his research dives in the early months of
1981 and 1982.
Gosliner took underwater photographs, and collected and preserved several
specimens (including CASIBP 034038 at the California Academy of Sciences, San
Francisco, examined by D. F. Dunn). His observations and comments are quoted
verbatim from letters:
‘We have been finding them commonly (about 5-6 individuals observed
during an hour dive). . . . Therefore I suspect that the populations are larger.
Generally specimens are loosely attached to the hydrocoral Allopora nobilis or
other arborescent organisms such as gorgonians. They are usually found on
vertical walls in areas of strong current, but do not appear to be orienting into the
current’ (17 February 1981).
Fig. 3. Specimen from
A NEW SEA ANEMONE 311
Oudekraal (SAM-—H2822, photos C. L. Griffiths). A. Approx. x 0,66.
Bo G2) CC Approx, x 12255 Dax I.
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fig. 4. A. Specimen from Oudekraal (SAM-—H2822, photo E. A. Robson). Vesicles have
shortened, revealing their irregular size and arrangement in relation to mesentery insertions.
Note flatter basal disc, and peristaltic wave starting below the oral disc. Scale in cm. B. Cnidae
from tentaculate vesicles (specimens SAM-—H3030): a, b—spirocyst at two levels of focus;
c, d, e—three basitrichs; f, g—an atrich at two levels of focus; h—atrich; i—discharged atrich;
j—discharged basitrich. Bar is 5 w. C. Preserved specimen of Capnella thyrsoidea from Hotten-
tot’s Huisie, Oudekraal (SAM-—H3169), x 1.
A NEW SEA ANEMONE 313
‘On a recent dive I observed that one specimen had completely engulfed all
but the basal stalk of an alcyonacean, Capnella [thyrsoidea]. The colony, which
was almost the size of the anemone, completely filled the gastrovascular cavity.
On a subsequent dive I observed three more anemones feeding on Capnella. One
specimen had half-digested a Capnella colony. An additional six specimens were
observed on that dive and all of them were within 10 cm of the remains of a basal
disc of a Capnella colony. From these observations I am entirely convinced that
[the] anemone is a species-specific predator on this alcyonacean. I think that the
presence of the anemones on vertical walls is not through accidental drifting but
through active association with their prey, which are most abundant on vertical
surfaces. Incidentally, I have never observed any drifting individuals. The animals
that I have seen have all been in 10-20 m of water. As Capnella occurs down to
about 150 [metres] I suspect that the anemones occur much deeper, as well.
.. . the strength of attachment I would equate . . . with that of a weakly attached
sea cucumber. I have never seen them use the pedal disc in their attachment.
I have only observed isolated individuals, never groups. I have seen them within
a foot of each other, however. . . . I have not observed any small individuals’
(14 April 1981).
‘The only thing I have seen the anemones “‘do” is eat or hang suspended by
their lateral tentacles. I am not sure how long feeding takes but as the whole
Capnella colony (except for the base) is digested including spicules, I would guess
it is a prolonged process. The pedal disc can be used to adhere to the substrate but
is always used in conjunction with the lateral tentacles. I have never observed
them adhering to the substrate solely by means of the pedal disc in “‘typical”’
anemone fashion’ (8 June 1982).
Most specimens seen by Gosliner were 10—20 cm in length. He estimates that
an expanded colony of Capnella thyrsoidea can reach 15 cm in height and perhaps
10 cm in diameter.
Photographs of specimens from Hottentot’s Huisie are shown in Figure
5A-D. The site is adjacent to Oudekraal and differs from it mainly in that there is
little kelp in the immediate vicinity owing to the steep topography and vertical
surfaces. The anemones seen here were all of similar colours. The specimen in
Figure 5A has the column and pedal disc pale scarlet, the vesicles iridescent white
with orange-red tips, and mesenterial insertions are marked as red lines. In
underwater photographs shown in Figure 5B—D, three anemones look brownish
pink, one paler than the others, the vesicles with tips of a darker colour, behind
which is often a white band. In one case (Fig. 5D) the tips of about ten vesicles,
greatly extended, are adhering to (probably) a Capnella colony to one side of the
anemone. This anemone is already attacking a colony situated in front of the oral
disc; exactly how it does this is not yet known.
Note on Capnella thyrsoidea (Alcyonacea, family Nephtheidae)
Capnella thyrsoidea (Verrill, 1865) is synonymous with C. gilchristi Thom-
son, 1910=C. rugosa Kikenthal, 1902 (Utinomi 1960; J. Verseveldt, pers.
314 ANNALS OF THE SOUTH AFRICAN MUSEUM
A NEW SEA ANEMONE 315
comm.). It appears to have a fairly wide distribution round the South African
coast at depths of 11-155 m, extending from the Atlantic coast of the Cape
Peninsula and False Bay to northern Natal (see Day et al. 1970; and the reference
collection of specimens in the Zoology Department, University of Cape Town,
identified by A. Tixier-Durivault). Kikenthal (1906) described C. rugosa from a
specimen obtained south of Mossel Bay at 155 m, and another was reported by
Thomson (1910) near East London at about 40 m. The usual substrate is probably
rock.
A specimen from Oudekraal collected and preserved by T. M. Gosliner is
shown in Figure 4C (SAM-H3169).
TAXONOMIC DESCRIPTION
(By K. W. England)
The discovery of the species of anemone described below necessitates
redefinition of the order Ptychodactiaria to which it belongs, and the creation of a
new family, the Preactiidae, to separate it from the other members of the order,
Ptychodactis patula Appellof (1893: 3-20) and Dactylanthus (Cystiactis) antarctica
(Clubb 1908: 5-6). These two species are contained in the family Ptychodac-
tiidae.
Order PTYCHODACTIARIA Carlgren, 1949 (redefined)
Diagnosis
Anthozoa (Dodecacorallia) with a definite base, with or without basilar
muscles; parietobasilar muscles present or absent. Filaments with or without
ciliated tracts. Gonads not enclosed in the mesogloea. Cnidome: spirocysts,
atrichs, heterotrichs and basitrichs.
Family Preactiidae fam. nov.
Diagnosis
Ptychodactiaria with column covered with tentaculate vesicles which may or
may not continue on to the periphery of the oral disc. Actinopharynx well
developed. Two siphonoglyphs supported by directive mesenteries. Twelve pairs
of mesenteries, mostly perfect, regularly arranged. All or almost all mesenteries
fertile. Filaments with ciliated tracts. Basilar muscles present.
Fig. 5 (facing page). Specimens of Preactis millardae sp. nov. from Hottentot’s Huisie, Oude-
kraal, photographed by T. M. Gosliner. Anemones seen there are 10 to 20 cm long. A. Living
specimen photographed in the laboratory. B—D. Underwater photographs of three individuals
in natural surroundings. B. Specimen attached to rock surface, stationary; oral disc to the right.
C. An expanded colony of Capnella thyrsoidea (lower right) being attacked. D. A third
individual feeding on C. thyrsoidea (to the right). Note that a few extended vesicles are attached
by their tips to another Capnella colony (upper left).
316 ANNALS OF THE SOUTH AFRICAN MUSEUM
Preactis gen. nov.
Etymology
As this genus has the characteristics of an actinarian on the one hand but
shows the characters of the Ptychodactiaria on the other, the name Preactis was
coined: pre from Latin prae = before or in advance, and actis from Actinia = sea
anemone, modern Latin from Greek axtic.
Diagnosis
Preactiidae without ectodermal longitudinal muscles on the column, except
where they continue for a short distance from the vesicles. Sphincter absent.
Actinopharynx with pocket-like prolongations and deep folds between the
siphonoglyphs. Siphonoglyphs well developed and considerably prolonged
beyond the length of the actinopharynx. The same number of mesenteries distally
and proximally. Mesenteries fused together axially for approximately one-third
‘the length of the column, from the base upwards. Cnidome: spirocysts, atrichs
and basitrichs.
Type species
Preactis millardae sp. nov.
Preactis millardae sp. nov.
Figs 2—4B, 5-10A
Material
The following description !s based on seven preserved specimens from three
localities. All were dissected and histological preparations were made as re-
quired.
Holotype
SAM-—H1677 in the South African Museum, Cape Town. Adult specimen
from rock pool at Onrus, near Hermanus, ccllected by M. R. Polchet, 4 February
IIE.
Paratypes
SAM-H2822 in the South African Museum, Cape Town, and BMNH
1983.4.27.1 in the British Museum (Natural History). Two large adult specimens
from Oudekraal, Cape Peninsula, depth 10-15 m, collected together by
C. L. Griffiths, 7 October 1976. CASIBP 034038 in California Academy of
Sciences, San Francisco. Specimen from Oudekraal, Cape Peninsula, depth 20 m,
collected by T. M. Gosliner, January 1981.
Additional material |
SAM-—H3030 in the South African Museum, Cape Town. Four small speci-
mens (three of these were examined here) from Maclear’s Beach, Cape Point
Nature Reserve, collected by S. X. Kannemeyer, 4 August 1980.
A NEW SEA ANEMONE 317
Sections of SAM-—H1677, SAM-—H2822, and BMNH 1983.4.27.1, by
K. W. England, in the British Museum (Natural History). Sections of CASIBP
034038, by D. F. Dunn, in the California Academy of Sciences, San Francisco.
Etymology
The species is named after Dr N. A. H. Millard, South African Museum,
Cape Town.
Description
Column conical, the base wider than the disc. The column completely
covered with tentaculate vesicles, arranged irregularly, though in places a vertical
or horizontal aignment may occur (see Fig. 2A—B). Near the margin the vesicles
occur singly or in pairs over both exo- and endocoels, but further down the
column the number in each exo- and endocoel may increase to three or four,
arranged in horizontal rows. The rows on the exocoels may or may not line up
with those on the adjacent endocoels, and vice versa (Fig. 4A). The vesicles are
conical with hemispherical tips, occasionally with a bifurcated tip. The uppermost
vesicles are positioned on or near the periphery of the oral disc and may encroach
on to the disc (Figs 2C, 3B). An occasional vesicle may occur on the base.
The disc is small and flat with a single central mouth.
The base is circular in outline, the diameter exceeding four times that of the
disc and capable of a much greater expansion. The mesenterial insertions can
easily be seen through the base tissue. The limbus is distinct.
Anatomy
The musculature of the column is weak and consists only of endodermal
circular muscle, which is only one fibre high (fibres less than 1 micron thick), and
the mesogloeal surface forms only simple folds (Fig. 8C). Sphincter absent.
Ectodermal longitudinal muscle fibres were not seen on the column, and if
present they would be less than 0,5 microns in diameter.
The tentaculate vesicles have both circular endodermal muscle and longitudi-
nal ectodermal muscle which continues for only a short distance on to the column.
Two distinct siphonoglyphs are present, supported by directive mesenteries,
and are prolonged aborally by as much again as the length of the actinopharynx.
Below the mouth the middle portion of the actinopharynx is greatly expanded and
in contraction it is thus thrown into several deep folds and pocket-like protuber-
ances in the area between the siphonoglyphs (Fig. 6). This arrangement increases
the potential diameter of the actinopharynx for feeding purposes.
The mesenteries are hexamerously arranged in two cycles, 6+ 6= 12 pairs.
In fully developed individuals all the mesenteries are perfect and fertile and bear
well-developed ciliated tracts in the uppermost region of the filaments. In younger
specimens some mesenteries of the second cycle may not be completely
developed. The lower parts of the perfect mesenteries are fused together axially,
the zone of fusion extending from the base upward for approximately one-third
318 ANNALS OF THE SOUTH AFRICAN MUSEUM
Disc
Actinophar ynx
Siphonoglyph
Ciliated tract
Main retractor
Secondary retractor
Gonad
Z = zone of fusion
Exocoelic muscle fibres
Fig. 6. Preactis millardae sp. nov. sectioned longitudinally with endocoelic view of one
mesentery (not to scale).
A NEW SEA ANEMONE 319
the length of the enteron. The fusion is not complete and openings along the line
of fusion remain, connecting the endo- and exocoels (Fig. 6).
The mesenteries possess large oral stomata, but marginal stomata are lacking.
The musculature of the mesentery is weak and unusual (Fig. 7). There are
two distinct retractor muscles. The main retractor extends from the base to the
discal region, some of the processes terminating under the disc but outer ones
ending on the column just below the disc. A secondary or pharyngeal retractor is
located behind the filaments, arising from the upper part of the fused portion of
the mesenteries and terminating in the region of the expanded middle portion of
the actinopharynx (Fig. 6). The reverse side of the mesentery lacks a
parietobasilar muscle and, instead, weak muscle bands arise at the top of the
fused portion of the mesenteries and fan out towards the column near the limbus,
but they become very diffuse and cannot be traced to the column wall (Fig. 6).
Very weak basilar muscles are present (Fig. 8A—B).
The gonads occur in a layer on either side of the mesogloea of the mesentery,
and they are not embedded in the mesogloea (Fig. 8D). They arise behind the
filaments, between the two retractors, and extend from below the actinopharynx
down to the region of the fused part of the mesenteries (Fig. 6).
Mesenteries
Inspection shows that mesenteries arise in the normal actinian sequence, and
this is reflected in the extent to which they are fused centrally. The first four
mesenteries formed—the first two couples—have the longest zones of fusion.
Shorter zones of fusion are found in turn in the two pairs of directive mesenteries
and in the last two couples, and the shortest are found in mesenteries of the
second cycle.
Mesenteries were fully formed in all the specimens examined except in three
instances, which seem to be second-cycle mesenteries in the course of
development. Their appearance suggests the following sequence of development,
although whether these examples are normal or exceptional cannot yet be
determined. A pair of mesenteries arises initially at the centre of the column as an
inverted pocket on the body wall, closed at the top and open at the bottom (see
Fig. 9, which corresponds to a dorsolateral exocoel in specimen BMNH
1983.4.27.1). The pocket grows upward and so moves towards the crown, while
splitting progressively along its midline so that two mesenteries are formed. These
also grow down the column towards the pedal disc, and filaments form on the free
edges below the pocket. Ciliated tracts develop in the appropriate region. It is
suggested that the two mesenteries become completely separated when the
pocket reaches the top of the column, after which they connect individually with
the oral disc and later the pharynx, and proximally with the pedal disc and with
the central region of mesenteric fusion.
In one specimen (from SAM-—H3030) a pocket giving rise to a pair of
second-cycle mesenteries in a lateral exocoel, the mesenteries showing filaments
with ciliated tracts and also retractor muscles, was almost fully enclosed in
320 ANNALS OF THE SOUTH AFRICAN MUSEUM
Filaments
Secondary
retractor =
Enlarged section of
main retractor
Main sretractor
OD.
ES SRS ama
mez ANU
Sms”
Fig. 7. Preactis millardae sp. nov. Transverse section of mesentery illustrating the position and
shape of the main and secondary retractors and the position of the gonad.
A NEW SEA ANEMONE 321
Fig. 8. Preactis millardae sp. nov. A. Basilar muscle in a vertical section of the pedal disc
(SAM-H1677). B. Basilar muscle in a similar section (SAM-—H2822). C. Circular muscle of
column in a vertical section below the margin (SAM-H1677). D. Section through the gonad,
male (SAM-—2822). Endoderm stippled, mesogloea white, ectoderm hatched.
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
Disc
Acti nopharynx
Column wall
Mesentery
Pocket of tissue
with developing
filaments
Filaments
Fig. 9. Preactis millardae sp. nov. Diagram of part of a specimen with pharynx and
disc cut open radially, showing the position of a pair of developing second-cycle
mesenteries: see text.
another pocket. The outer pocket, very large, extended from below the oral disc
downward to further even than mid-column. It consisted of two first-cycle
mesenteries still joined for two-thirds of their length, the pocket having failed to
split along its midline. There were thus no filaments although retractor mucles
and gonads were present. These first-cycle mesenteries were also anomalous in
being from adjacent pairs (the ventro- and dorsolaterals).
Cnidome
The size ranges and types of cnidae observed are given in Table 1, and the
types illustrated in Figures 4B and 10A. Of particular interest is the spirocyst in
which the helical folds of the undischarged thread can be seen with the light
microscope (Fig. 4B, a—b).
A NEW SEA ANEMONE 323
TABLE 1
Cnidae of Preactis millardae gen. et sp. nov. (measurements in microns).
Location/Type SAM-H1677 SAM-H2822 BMNH 1983.4.27.1
Tentacles
Spirocysts 13,8-16,3 x 2,5 14,6-17,9 x 2,0 14,6-20,0 x 2,6-3,3
Basitrichs 12,5-16,3 X2,5-3,0} 15,2-21,2 (23,8) x2,0-3,3 | 13,3-17,9 x 2,0-2,6
Atrichs 20,0—26,3 x 2,5 20,5—27,8 X 2,6—4,0 17,9-31,2 x 2,6—4,0
Disc
Basitrichs 13,3-15,2 x 2,0-2,6 13,3-17,2 x 2,0-2,6
Pharynx
Atrichs 20,0—22,5 x 2,5-3,0] 17,2—23,2 x 2,0-3,3 15,2-25,2 x 2,0-3,3
Basitrichs — 16,6 x 2,0 (1 only) 15,9-17,9 x 2,6 (3 only)
Filaments
Atrichs 16,3—22,5 x 2,5 17,9-24,5 (27,8) x 2,0-3,3
Basitrichs — 13,3-15,9 x 2,0 (few)
The size ranges were established by measuring the smallest and largest of
each type found, from at least three separate squash preparations taken from
different sites of each area examined, 1.e., the tentacles, disc, actinopharynx, and
filaments. Examination was carried out using normal light microscopy (bright
field and phase contrast).
Identification
From the foregoing description it is evident that this species resembles in
many respects Dactylanthus antarcticus Carlgren, 1911 (Dunn 1983). The possibil-
ity of Dactylanthus being a young or less well-developed form of the present
species was considered, and so the specimen of D. antarcticus described and
identified by Stephenson (1918) from the Terra Nova Expedition was examined
(BMNH 1918.5.12.4).
Although histological sections were not made, a few new details can be
added to the descriptions given by Clubb (1908) and Carlgren (1911). Dactylan-
thus has a double retractor muscle similar to that of the new species, the second or
pharyngeal retractor being in the same position. The gonads similarly occur
between the two retractors but they are lower down the column than in the new
species, although not completely below the filaments as stated and illustrated by
Carlgren (1911, pl. 1 (fig. 3)). The specimen shows no evidence of parietobasilar
muscles.
The nematocysts of Dactylanthyus antarcticus were examined and good
agreement with the size ranges given by Carlgren (1940) was achieved. Dunn
(1983) redescribed D. antarcticus and her measurements for the nematocysts are
slightly larger than here but of the same order of magnitude. In addition, hetero-
trichs were found and a few nematocysts were seen that resembled holotrichs. It is
thought that the latter were probably developing atrichs, but further work is
required to substantiate this possibility. The types and size ranges of cnidae found
were as follows:
324 ANNALS OF THE SOUTH AFRICAN MUSEUM
Vesicles of column: Spirocysts 16,3—21,3 X approx. 2,0
Atrichs 18,8-31,7 x 3,0-4,0 wu
Heterotrichs 7,5—-15,6 X approx. 2,0 u
Actinopharynx: Atrichs 11,3-23,2 x 2,0-3,3 uw
Filaments: Atrichs 11,3-18,8 x 2,0-2,5 yu
These are illustrated in Figure 10B. The differences between the nematocysts
of Dactylanthus and of the new species can be seen by comparing the cnidae of
Figure 10A-B. Although the types of nematocysts are the same in both, the
spirocysts differ: in Dactylanthus they are more like those found in the Actiniaria
in that the folds of the thread are not visible and the thread occupies more of the
capsule.
From the study of cnidae it is clear that Dactylanthus is not a young form of
the new species.
The taxonomic characters of the new species and those of Dactylanthus are
given in Table 2, listed in the order in which they would be examined, together
with those of Ptychodactis patula Appellof, 1893. It can now be seen that the new
species differs from both the others in three features that are of major taxonomic
importance, i.e.
(1) ectodermal longitudinal muscle of the column is absent,
(ii) ciliated tracts are present on all filaments,
(iii) basilar muscles are present.
TABLE 2
Comparison of the major characters of Ptychodactis, Dactylanthus, and
Preactis gen. nov.
Column smooth.
Tentacles present on oral disc.
Ectodermal longitudinal muscle
present on column.
No sphincter.
Up to four cycles of mesenteries
present.
Mesenteries not fused together
near the base.
No ciliated tracts.
Gonads not enclosed in the me-
sogloea.
Basilar muscles absent.
Parietobasilar muscles present.
Column with single vertical rows
of tentaculate vesicles over each
endo- and exocoel.
No tentacles on oral disc but
peripheral tentaculate vesicles.
Ectodermal longitudinal muscle
present on column.
Weak sphincter present.
Six pairs of mesenteries perfect.
Mesenteries fused together
axially near the base.
No ciliated tracts, but filaments
of second cycle terminate in bi-
lobed funnel.
Gonads not enclosed in the me-
sogloea.
Basilar muscles absent.
Parietobasilar muscles absent
(p.323).
Column with multiple irregular
rows of tentaculate vesicles over
each endo- and exocoel.
No tentacles on oral disc but
peripheral tentaculate vesicles.
No ectodermal longitudinal mus-
cle on column.
No sphincter.
Twelve pairs of mesenteries per-
fect.
Mesenteries fused together
axially near the base.
Ciliated tracts present, bilobed
funnels absent.
Gonads not enclosed in the me-
sogloea.
Basilar muscles present.
Parietobasilar muscles absent.
A NEW SEA ANEMONE 325
‘S
oO
2 WEK*KKKKKE
4)
Ss
0 SSS
Z
A B
a 2
yy =
Zz, JU
| y
] —_z Si
Z —Z Zy
y SF =]
Z = Z
7, =
ae) Ss
y, SZ Cc
~
Ss,
SS
SI
b
B
Fig. 10. A. Preactis millardae sp. nov. Cnidae (see Table 1). Vesicles: a—spirocyst, b—basitrich,
c—atrich. Pharynx: d—atrich, e—basitrich. Filaments: f—atrich, g—basitrich.
B. Dactylanthus antarcticus (BMNH 1918.5.12.4). Cnidae. Vesicles: a—spirocyst, b—atrich,
c—heterotrich. Pharynx: d—atrich. Filaments: e—atrich.
These features prevent the new species from being referred either to Dacty-
lanthus or to Ptychodactis. Furthermore, they would even rule out the inclusion of
this species in the Ptychodactiaria as defined by Carlgren (1949) and would
suggest that it belongs to the Actiniaria. However, as the gonads are not enclosed
in the mesogloea of the mesenteries, the Actiniaria are precluded.
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
The combination of these few but significant features with gonads not con-
tained in the mesogloea suggests a species that in some respects may be interme-
diate between the Ptychodactiaria and the Actiniaria, and the name Preactis gen.
nov. is proposed to reflect this.
It is also proposed to redefine the order Ptychodactiaria to accommodate the
new genus (p. 315). The primary character of the Ptychodactiaria becomes the
presence of gonads not enclosed in the mesogloea, and other characters in
Carlgren’s (1949) diagnosis are modified, i.e. ciliated tracts present or absent, and
basilar muscles present or absent.
If Carlgren’s taxonomic criteria are applied it is not possible to include
Preactis in the family Ptychodactiidae. Carlgren (1942) considered the presence
or absence of basilar muscles an important character of fairly high classificatory
value, and he did not place species with and without them in one family. The
presence or absence of ciliated tracts on the filaments is of similar significance. In
order to accommodate Preactis, which possesses basilar muscles and ciliated
tracts (whereas Dactylanthus and Ptychodactis do not), a new family in the
Ptychodactiaria is proposed. The diagnosis of this family, the Preactidae fam.
nov., is given on p. 315.
Order Ptychodactiaria
Ptychodactiidae Preactiidae
genus Ptychodactis genus Preaciis
genus Dactylanthus
It is clear that further research is needed to resolve the taxonomic anomalies
offered by the Ptychodactiidae, for it will be appreciated that Dactylanthus and
Preactis are much more alike than Dactylanthus and Ptychodactis.
COMMENTS
The affinities of the Ptychodactiaria and their geographical distribution
deserve further thought.
The higher taxa of Anthozoa as Carlgren saw them in 1949 are these:
ANTHOZOA
Subclass Zoantharia
Orders Ptychodactiaria
Corallimorpharia
Actiniaria
Madreporaria
Zoantharia
Antipatharia
Ceriantharia
Subclass Alcyonaria
A NEW SEA ANEMONE S27
In 1944 he expressed his views on ‘the lines of development of the Anthozoa’
as a diagram (see, however, the more recent views of Schmidt & Zissler 1979).
Carlgren’s graphic presentation is conveyed in Figure 11. He considered the
Ptychodactiaria as ‘a primitive group which has separated early from the Acti-
niaria and gone their own way’ (Carlgren 1942).
Actiniaria
c Antipatharia
= 2
@ E orena ©
o ; orallim . o
(S) (6)
91 Mag =
6| x—"2Pora ria oanth A
a] S xr
a »
Rugesa :
Ceriantharia
tych Odac tiarja
@
8
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O /
e) j
Fig. 11. Diagram after Carlgren (1944), depicting his view of lines of
development in the Anthozoa.
Whether the Ptychodactiaria are primitive or, as judged earlier by Stephen-
son (1921), ‘a collection of curiosities which have developed along a little line of
their own’, new observations are now needed on Ptychodactis patula, if possible
on living material. It has an arctic-boreal distribution (Carlgren 1949) and Appel-
16f’s (1893) specimens were found on gorgonians. Details of its biology are quite
unknown, however, and the same is true of Dactylanthus. The functions of special
anatomical features are difficult to infer from preserved specimens and the
muscular and ciliary systems, for example, need re-examination.
The features of Dactylanthus that separate it from Ptychodactis and relate it
to Preactis may be considered to include its Antarctic distribution. The distribu-
tion of D. antarcticus is summarized by Dunn (1983) as south of the 64th parallel,
328 ANNALS OF THE SOUTH AFRICAN MUSEUM
on both sides of Drake Passage between 56° and 65°W, and in the Ross Sea and
north of it (162°W to 170°E). A circumpolar distribution of this kind would have
followed the isolation of the Antarctic continent (i.e. post-Eocene: Frakes 1979).
It is possible to speculate that if Preactis were derived from the same ancestral
Antarctic stock as Dactylanthus the formerly narrower polar seas might have
allowed ancestral pelagic larval stages to be carried northward by currents to-
wards southern Africa and elsewhere. Although the life history of Ptychodactiaria
is still unknown, Dactylanthus, Preactis and Ptychodactis all have numerous and
small oocytes and the existence of pelagic larvae seems probable.
In view of the distribution of Preactis millardae (Fig. 1) it is worth mentioning
that Carlgren (1938) thought two species of littoral sea anemones from the
Atlantic coast of the Cape Peninsula to be ‘certainly of subantarctic origin’.
Phellia aucklandica, collected from Oudekraal at low water by T. A. Stephenson,
he found identical to specimens from the Auckland Islands, the Inaccessible
- Archipelago and Gough Island. Helianthella annularis, from Oudekraal and
Kommetjie, is an intertidal species with a brood pouch first described by Carlgren
(1938). He considered it to show a close resemblance to H. kerguelensis found at
Kerguelen and Macquarie Island.
The discovery of Preactis millardae raises very interesting questions on the
taxonomy, zoogeographical history and phylogeny of the Ptychodactiaria and it is
hoped that a greater awareness of these animals may lead to further work on their
biology.
ACKNOWLEDGEMENTS
Several colleagues have contributed materially to this paper: without Pro-
fessor J. H. Day, Dr T. M. Gosliner, Dr C. L. Griffiths and Dr N. A. H. Millard
there would be few recorded observations and fewer, if any, specimens. It is a
pleasure to thank them for their generous help. Dr R. J. Griffiths, M. R. Polchet
and Mr S. X. Kannemeyer are thanked for collecting some of the specimens. A
visit by E. A. Robson to the Zoology Department, University of Cape Town, in
1976, for which thanks are due to Professor A. C. Brown, made it possible
subsequently to collate existing records and to include them here. Dr D. F. Dunn
is thanked for valuable comments and for sending us her review of Dactylanthus
before publication. Dr J. Verseveldt of the Rijksmuseum van Natuurlijke His-
torie, Leiden, kindly identified the specimen of Capnella thyrsoidea shown in
Figure 4C.
NOTE IN PRESS
The authors have seen photographs taken by S. Gerber of a probable further specimen of
this anemone. The specimen was observed near Gordon’s Bay, nearly due east from Simonstown
at the opposite side of False Bay (April 1984, 25 m depth, 5,7 km offshore). We wish to thank Mr
S. Gerber from Stellenbosch and Miss E. Louw of the South African Museum for providing this
information.
A NEW SEA ANEMONE 329
REERERENCES
ANDREWS, W. R. H. & Hurcuines, L. 1980. Upwelling in the Southern Benguela Current.
Progr. Oceanogr. 9: 1-81.
APPELLOF, A. 1893. Ptychodactis patula n.g. und sp. der Representant einer neuen Hexactinien
Familie. Bergens Mus. Arb. 4: 1-22.
Brown, A. C. & JARMAN. N. 1978. Coastal marine habitats. Jn: WERGNER, M. J. A. ed.
Biogeography and ecology of southern Africa. The Hague: Junk. Monographiae Biol. 31:
1239-1277.
CARLGREN, O. 1911. Uber Dactylarthus (Cystiactis) antarcticus (Clubb). Wiss. Ergebn. schwed.
Stidpolarexped. 6(5): 1-31.
CARLGREN, O. 1938. South African Actiniaria and Zoantharia. K. svenska VetenskAkad. Hand.
(3) 17(3): 1-148.
CARLGREN, O. 1940. A contribution to the knowledge of the structure of the cnidae in the
Anthozoa. Lunds Univ. Arsskrift. N.F. Avd 2 36(3): 1-62.
CaRLGREN, O. 1942. Actiniaria II. Dan. Ingolf-Exped. 5(12): 1-92.
CARLGREN, O. 1944. Das System und die Entwickiungslinien der Anthozoen zugleich einige
Bemerkungen iiber Pax’ Bearbeitung dieser Tiergruppe in der “Tierwelt der Nord- und
Osisee’. K. fysiogr. Sdllsk. Lund Férh. 14(4): 40-SS.
CaRLGREN, O. 1949. A survey of the Ptychodactiaria, Corallimorpharia and Actiniaria.
K. svenska VetenskAkad. Handl. (4) 1(1): 1-121.
Cus, J. A. 1908. Coelentera IV. Actiniae. Nat. Antarct. Exped. 1901-1904. Nat. Hist. 4(9):
1-12.
Day, J. H. 1970. The biology of False Bay, South Africa. Trans. R. Soc. S. Afr. 39: 211-221.
Day, J. H., Fretp, J. G. & PenritH, M. J. 1970. The benthic fauna and fishes of False Bay,
South Africa. Trans. R. Soc. S. Afr. 39: 1-108.
Dunn, D. F. 1983. Some Antarctic and Sub-Antarctic sea anemones (Coelenterata: Ptychodac-
tiaria and Actiniaria). Jn: KoRNICKER, L. S. ed. Biology of the Antarctic seas XIV. Antarctic
Res. Ser. Washington. 39: 1-67.
FIELD, J. G., Grirfirus, C. L., LINLEY, E. A., CARTER, R. A. & ZOUTENDYK, P. 1980.
Upwelling in a nearshore marine ecosystem and its biological implications. Estuar. cst. mar.
Sci. 11: 133-150.
Frakes, L. A. 1979. Climates through geologic time. Amsterdam: Elsevier.
KUKENTHAL, W. von. 1902. Diagnosen neuer Alcyonarien aus der Ausbeute der Deutschen
Tiefsee-Expedition. Zool. Anz. 25: 299-303.
KUKENTHAL, W. von. 1906. Alcyonacea. Wiss. Ergebn. dt. Tiefsee-Exped. ‘Valdivia’ 13(1) Lief.
ett.
ScHmipT, H. & ZIssLER, D. 1979. Die Spermien der Anthozoen und ihrer phylogenetische
Bedeutung. Zoologica, Stuttg. 44(129): 1-92.
STEPHENSON, T. A. 1918. Coelenterata. I. Actiniaria. Br. Antarct. Terra Nova Exped. 1910.
(Zool.) 5 (1): 1-68.
STEPHENSON, T. A. 1921. On the classification of Actiniaria. II. Considerations of the whole
group and its relationships, with special reference to forms not treated in Part I. Quart. J.
micr. Sci. 65: 493-576.
Tuomson, J. S. 1910. The Alcyonaria of the Cape of Good Hope and Natal. Trans. R. Soc.
Edinb. 47: 549-589.
Utinomi, H. 1960. A revision of the nomenclature of the family Nephtheidae (Octocorallia:
Alcyonacea). I. The genera Capnella, Scleronephthya and Chondronephthya (n.g.). Publs
Seto mar. biol. Lab. 8: 27-40.
VELIMIROV, B., FIELD, J. G., GriFFITHS, C. L. & ZOUTENDYK, P. 1977. The ecology of kelp bed
communities in the Benguela upwelling system. Helgoldnder wiss. Meeresunters. 30:
495-518.
VERRILL, A. E. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedi-
tion, under Commodore C. Ringgold and Captain John Rodgers, U.S.N., from 1853 to
1856. Part II. Alcyonaria. Proc. Essex Inst. 4(12): 181-196.
6. SYSTEMATIC papers must conform to the /nternational code of zoological nomenclature
(particularly Articles 22 and 51).
Names of new taxa, combinations, synonyms, etc., when used for the first time, must be
followed by the appropriate Latin (not English) abbreviation, e.g. gen. nov., sp. nov., comb.
nov., syn. nov., etc.
An author’s name when cited must follow the name of the taxon without intervening
punctuation and not be abbreviated; if the year is added, a comma must separate author’s
name and year. The author’s name (and date, if cited) must be placed in parentheses if a
species or subspecies is transferred from its original genus. The name of a subsequent user of
a scientific name must be separated from the scientific name by a colon.
Synonymy arrangement should be according to chronology of names, i.e. all published
scientific names by which the species previously has been designated are listed in chronological
order, with all references to that name following in chronological order, e.g.:
Family Nuculanidae
Nuculana (Lembulus) bicuspidata (Gould, 1845)
: Figs 14-15A
Nucula (Leda) bicuspidata Gould, 1845: 37.
Leda plicifera A. Adams, 1856: 50.
Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (fig. 8a—b).
Nucula largillierti Philippi, 1861: 87.
Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.
Note punctuation in the above example:
comma separates author’s name and year
semicolon separates more than one reference by the same author
full stop separates references by different authors
figures of plates are enclosed in parentheses to distinguish them from text-figures
dash, not comma, separates consecutive numbers
Synonymy arrangement according to chronology of bibliographic references, whereby
the year is placed in front of each entry, and the synonym repeated in full for each entry, is
not acceptable.
In describing new Species, one specimen must be designated as the holotype; other speci-
mens mentioned in the original description are to be designated paratypes; additional material
not regarded as paratypes should be listed separately. The complete data (registration number,
depository, description of specimen, locality, collector, date) of the holotype and paratypes
must be recorded, e.g.:
Holotype
SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach
Port Elizabeth (33°51’S 25°39’E), collected by A. Smith, 15 January 1973.
Note standard form of writing South African Museum registration numbers and date.
7. SPECIAL HOUSE RULES
Capital initial letters
(a) The Figures, Maps and Tables of the paper when referred to in the text
e.g. ‘... the Figure depicting C. namacolus ...’; ‘. . . in C. namacolus (Fig. 10)...’
(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded
by initials or full names
e.g. Du Toit but A.L.du Toit; Von Huene but F. von Huene
(c) Scientific names, but not their vernacular derivatives
e.g. Therocephalia, but therocephalian
Punctuation should be loose, omitting all not strictly necessary
Reference to the author should be expressed in the third person
Roman numerals should be converted to arabic, except when forming part of the title of a
book or article, such as
‘Revision of the Crustacea. Part VIII. The Amphipoda.’
Specific name must not stand alone, but be preceded by the generic name or its abbreviation
to initial capital letter, provided the same generic name is used consecutively.
Name of new genus or species is not to be included in the title: it should be included in the
abstract, counter to Recommendation 23 of the Code, to meet the requirements of
Biological Abstracts.
K. W. ENGLAND & E. A. ROBSON
A NEW SEA ANEMONE FROM SOUTH AFRICA
(ANTHOZOA, PTYCHODACTIARIA)
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