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VOLUME XLV
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1959 — 1960
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LIST OF CONTRIBUTORS
BARNARD, K. H.
Contributions to the knowledge of South African marine mollusca. Part II.
Gastropoda: Prosobranchiata: Rhachiglossa (published June 1959) ..
Bone, E. L.
See SINGER, R.
Brown, A. C.
A new Solifugid Arachnid from Table ee oe es eee sp. n.
(published July 1960) : :
DAY, J. H.
The Polychaet Fauna of South Africa. Part 5. Errant species eae ecg off sa
Coasts (published May 1960)
Juss, R. A.
. Note on locality records of Freshwater Fishes presented by F. D. McKean to the
South African Museum (published April 1960) .
Minrarp, N. A. H.
Hydrozoa from ships’ hulls and eee i in ite Town Docks ae
July 1959)
Sincer, R., and Bonk, E. L.
Modern Giraffes and the Fossil Giraffids of Africa (published July 1960)
Warnor, F. H.
Additions to the South African Museum Collection of Marine Fishes eee
April 1960) : = - ae as é sae
Ta.sor, F. H.
Notes on the Biology of the Lutjanidae (Pisces) of the East African Coast, with
special reference to L. bohar (Forskal) (published July 1960)
SMii }HSOT NUAN
INSTITUTION
Page
DD
261
260
239
375
257
549
JUN 7 1961
‘i
oa
ALL
i.
ae iy
We. fo
IN THIS VOLUME
. : Page
s (Eunicidae), Day, 1960 va oy: oe Sy, ae Be iar vio)
syllis (Syllidae), Day, 1960... ne eg ite ot as be Be)
Adinopsis
Afritrophon
Afrivoluta
Afrocominella
Aglaophamus
Allothunnus ..
Amblyosyllis ..
Anaitis ..
Anaitides
Ancilla
Antinoe
Aphareus
Aphrodita
Aprion
Arabella
_ Aspella
Autolytus
Babylonia
Barbourisia
Bhawania
Bougainvillia ..
Bramatherium
Bullia ..
Burnupena
Callithea
Campanularia ..
Cancellaria
Cantharus
Charitodoron..
Chloeia
Clavisyllis
Clytia ..
Columbarium
Columbella
Cominella
Coralliobia
Coralliophila ..
Cronia
Ctenopoma
Demoulia
Dibaphus
LIST OF GENERA AND SUBGENERA
Page
142
207
23
152
327
258
313
208
296
61
281
568
274
551
363
232
317
147
257
294
242
519
123
158
50
248
12
150
i44
295
320
248
234
180
169
193
189
229
260
122
RE
Diopatra
Dorsanum
Dorvillea
Drilognathus ..
Drilonereis
Drupa..
Drupella
Ehlersia
Engina
Enipio. .
Ephesia
Epidiopatra
Epinephelus ..
Eteone
Eulagisca
Eulalia
Eumida
Eunice
Euphione
Euphrosyne
Eurythoe
Euthalenessa ..
Euthria
Exogone
Fasciolaria
Fulgoraria
Fusivoluta
Fusus ..
Galeodes
Genetyllis
Giraffa
Giraffokeryx ..
Clycera
Glycinde
Goniada
Gonothyraea
Griquatherium
Grubea
Halosydna
Haplochromis
466 et passim
143
298
375 et passim
519
328
331
333
249
315
282
260
Haplosyllis
Harmothoe
Harpa.. he
Helladotherium
Hemilepidia
‘Hindsia
Hololepida
Honanotherium
Hyalinoecia
Hydaspetherium
Hypoeulalia
Imbricaria
Indratherium
Kefersteinia
Kirchenpaueria
Labrorostratus
Laeonereis
Lagisca Ae
Lamellisyllis ..
Langerhansia
Laomedea
Latiaxis
Latirus
Leiodomus
Leodice .. f
Lepidasthenia
Lepidonotus ..
Leptoecia
Libytherium ..
Lovenella
Lumbrineris ..
Lutjanus
Lysidice
Malmgrenia .
Marginella
Marphysa
Mastigonereis ..
Melapium
Melongena ied
Micronephthys ..
Mitra .
Murex..
Myrianida
Mysta ..
Nassa ..
Nassaria
Neanthes
Nephthys
1A
LIST OF GENERA AND SUBGENERA
498,
-- 491,495,
Neptuneopsis . .
Nereis...
Nothria
Notocirrus ;
Notophyllum ..
Stauronereis
Obelia. . ae
Odontosyllis ..
Okapia
Oliva .
Onuphis 2
Ophioglycera..
Orangiatherium
Orasius
Orthopyxis
Palaeotragus ..
Paleanotus
Pareurythoe
Pelmatochromis
Perigonimus
Perinereis
Peristernia
Pharyngeovalvata
Pholoe. .
Phyllodoce
Phyllosyllis
Pionosyllis
Platynereis
Plumularia
Polyeunoa
Polynoe
Pristipomoides
Procerastea
Progirafia
Propalaeomeryx
Protomystides
Psammolyce ..
Pseudonereis ..
Pterocirrus
Pterosyllis
Pterothrissus ..
Pusia ..
Pusiostoma
Pyrene
Rapana ;
Rhamphobrachium ..
Rhizorhagium
Samotherium. .
Sarsia ..
3 330
-+ 447,491, 525
515, 519
248
515, 519
294,
295
LIST OF GENERA AND SUBGENERA
Page Page
281 ritonalia) 4 52 Bs re sta wig 212
260 Trophon an we wis set Od
290 Trypanosyllis . . ee aed eh wees
303 Tubulara -. :. oe ae sewn, 240
417 et passim Turricula a he we Se 52
a 8595 Typhis es a ks Sig J 2TO
ee 320
- Sphaerosyllis . . 316 U
Staurocephalus .. 371 Urosalpinx .. as G te ragt
Stauronereis .. ee Sees < .
*Steggoa.. a es e Sur GOO My
_ Stegolophodon ae Bs Se. RBA Vasum a vi Re os 36
-Sthenelais .. ia ae 25 |: 289 Vexillum as aye ae ae 52
_ Syllides oe i ue se REO Vishnutherium ans as ye ARNO
- Syllidia ss ue wa al ee SOm Volema bs be afi gel an TAS
“Syllis .. me oY ia ee SOT Voluta Ms ze as M, 18
Sylvanocochlis ue ‘ As 60 Volutocorbis .. “es ss, ae 24
; Syncoryne <a se - ee ==
T Xancus cs ae br 28 36
PePhais .. = ae siete Maar
Thalenessa .. See bie et) 204 Z
_ Tilapia oye sof af eee 260 Zemiropsis .. ne be e200, 147
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CONTRIBUTIONS TO THE KNOWLEDGE OF SOUTH AFRICAN
MARINE MOLLUSCA. PART II. GASTROPODA:
PROSOBRANCHIATA: RHACHIGLOSSA*
By K. H. BARNARD
(With 52 figures in the text)
RHACHIGLOSSA
Fam. MARGINELLIDAE
1917. Tomlin. Proc. Mal. Soc., xii, pp. 242-306 (list of Recent species).
1925. Thiele. D. Tiefsee Exp., xvii, pp. 191-7.
1932. Turton. Mar. Sh. Port Alfred, pp. 33-44.
A large number of specimens, fresh and worn, have been retrieved from
the P.F. bottom-samples. Most of these have been identified; the remainder
may be useful when a thorough study of South African Marginellas is under-
taken.
The Pieter Faure did not obtain all the species described by Thiele from the
Valdivia collection.
In this work only those species are mentioned for which additional
records extend the hitherto known distribution.
Tomlin (p. 246) credits 43 species to the ‘Cape’ (South African) region.
In later years he added four more species. All of these seem to be weil estab-
lished, though I have suggested (znfra) that taylori renamed barnardi is a synonym
of differens.
Thiele added 12 species, but I cannot admit the validity of all of these.
Turton listed 79 names of ‘species and varieties’ found by him at Port
Alfred. Many of these are obviously additional names applied to known
species and juveniles.
About 60 species might be a fair estimate of the number of South African
Marginellas.
The size of apparently mature examples of the same species sometimes
varies considerably: e.g. musica (with fully developed outer lip) from 15 to
25 mm. Tomlin has no comment on this feature, but he mentions that sinis-
trorsity occurs more frequently in this family than in other marine Gastropods
(p. 246). I have not seen one such example, though some species (e.g.
biannulata) are often found in hundreds in private collections.
* Part I. Toxoglossa: Ann. S. Afr. Mus., vol. xliv, part 4, 1958.
I
VOL. XLV PART I
TETRA aug x4 fe’
2 ANNALS OF THE SOUTH AFRICAN MUSEUM
The presence or absence of denticles (plicae) within the outer lip is
usually easily seen, but the lip margin is much exposed to wear and often appears
smooth. This may explain Smith’s statement that the lip is either smooth or
minutely denticulate in dulcis (= bensont).
The radulae of only two species have been obtained: capensis and biannu-
lata. There was no difficulty in finding the radulae of these two; but examples
of rosea, bairstowi and musica were examined without success. Evidently the
radulae of these species are very small and only to be found by a more refined
technique than is usually employed. Tomlin (p. 245) quotes Gwatkin as
saying that the radula of Marginellas is ‘extremely hard to extract’ but whether
the difficulty is due to minute size is not stated. Fresh material is necessary, as
animals long preserved in formalin or alcohol are not easily dissected.
For ‘Marginella’ angustata Sow. (Smith, 1906, Ann. Natal Mus., 1, p. 28),
see Ancilla errorum ‘Tomlin.
Egg-capsules. ‘Twelve egg-capsules affixed to a Clavatula tripartita from
31° 38’ S. 29° 34’ E. (off Port St. Johns), 26 fathoms (U'C igre
Nearly hemispherical, maj. and min. diam. 4 and 3, height 2 mm.
Only one protoconch in each capsule. 24 whorls, alt. 2:5, diam. 2 mm.
White, with on the last half or three-quarter whorl a blackish-grey band, more
or less broken up into spots, at the suture with previous whorl. Columella
pleats 4.
A 45 mm. nebulosa Bolten (S. Afr. Mus. no. A6382) has a similar dark
stripe on the Ist postnatal whorl, but not on the protoconch (if there was one,
it has been obscured by the surface glaze).
The protoconchs of bicatenata and mosaica also seem to be of a suitable size,
but the juveniles (v. infra, pp. 5, 6) show no trace of a dark band at the top of
the whorl near the suture.
M. piperata also is not excluded as a possible parent of these capsules, but
no unworn examples are available.
Marginella munda Smith
1904. Smith. 7. Malac., xi, p. 31, pl. 2, fig. 14.
Off Illovo River (Natal), 27-30 fathoms, one worn and broken; off Cove
Rock (East London area), 22 fathoms, two worn (S. Afr. Mus. P.F. coll.).
Marginella aphanospira Tomlin
1913. Tomlin. 7. Conch., xiv, p. 101 text-fig.
The type was described as having two columella pleats. The specimen
seen and referred to by Tomlin (p. 102) has 3 pleats. The 3rd pleat can only
be seen when the shell is viewed obliquely from below (except in immature
examples where it is fully exposed). A Still Bay example has 3 and a feeble
4th pleat.
Port Elizabeth (seen by Tomlin) and Still Bay (S. Afr. Mus.).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 3
Off Umkomaas River (Natal), 40 fathoms, 2; off Sandy Point (north of
Cape Morgan), 51 fathoms, 2; off Cape Morgan, 33 fathoms, one; off Hood
Point (East London area), 49 fathoms, 2; 34° 5’ S., 25° 55’ E., 67 fathoms, one
femiature;, 394° 5. 25° 44 E., depth ?, 2 (S. Afr. Mus. P.F..coll.).
Marginella capensis Krauss
q Fig. 1(d)
1848. Krauss. Siidafrik. Moll., p. 25, pl. 6, fig. 21.
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), pp. 34, 49.
When collected alive the shell is fawn coloured, the sutures marked by a
narrow white band, outer lip and anterior portion of the columella white.
Radula with 46-56 rows, the denticles variable and asymmetrical on
either side of the median line: see fig. 1d.
Seven specimens from 34° 27’ S. 25° 42’ E., 256 fathoms, were identified
___ by Sowerby as this species, although the largest one measures only 9 X 4:3 mm.
7 In shape they resemble fallax more than typical capensis, but there are only
- 4 columella pleats and no trace of denticulations within the outer lip. They
probably represent another species, but for the present may be recorded here.
Marginella perla Marrat
Fig. 1(a)
1852. Krauss. Arch. Naturg., i, p. 37 (biplicata, preocc.).
1876. Marrat. 7. Conch., i, p. 136.
1886. Watson. Challenger Rep., xv, p. 267, pl. 16, fig. 8 (chrysea).
1903.* Von Martens. D. Tiefsee Exp., vii, pl. 3, fig. 6 (Krauss’s type) (not the recorded
: specimen = brocktonz).
1932. Turton. loc. cit., p. 36, pl. 7, no. 272 (2nnocens).
A curious feature of this species is the retention of previous lip varices on
the outer surface. This is referred to by Watson, but not shown in his figure.
It appears, however, in von Martens’s figure of Krauss’s type. It occurs in all
the specimens I have seen, and may be regarded as a diagnostic character.
Sometimes it gives an almost turreted appearance to the spire.
Watson gave the number of columella pleats as 3, but the 3rd (uppermost)
is scarcely a pleat: ‘only the end of the columella surface before the second
pleat’ (von Martens).
Sea Point, Cape Town (Watson); St. James (False Bay), and Still Bay
(S. Afr. Mus.); Port Elizabeth (Sowerby); Port Alfred (Turton).
* In Part I the date of this paper was quoted as 1904, because it was received at the British
| Museum Library in February 1904, and was also noticed in Naturae Novitates in the same month.
I have since found, however, that the original cover of Lief. 1 of vol. vii is dated 1903. Von
: Martens’s paper is also entered in the 1903 Zoological Record.
‘, The exact month of publication of von Martens’s paper is sometimes important to decide
priority, e.g. Nassa analogica Sow., 8 July 1903, N. circumtexta von Martens, 18 Dec. 1903.
4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Marginella lucida Marrat
1877. Marrat. 7. Conch., 1, p. 205.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p- 39, pl. 1, fig. 2 (turtont).
Off Tugela River (Natal-Zululand), 47 fathoms, one; off Illovo River
(Natal), 27-30 fathoms, 3; off Umkomaas River (Natal), 40 fathoms, one;
off Cape Morgan, 47 fathoms, 4; off East London, 20 fathoms, one; off Cove
Rock (East London area), 22 fathoms, 3 (S. Afr. Mus. P.F. coll.). g0° 47'S.
30° 29’ E., 24 fathoms (U.C.T.).
a da
Fie; 17,
(a) Marginella perla Marrat. (b) M. perminima Sow. (c) radula plate of M. biannulata (Fabr.).
(d) radula plates from two individuals of M. capensis Krss. to show variation.
Marginella biannulata (Fabr.)
Fig. 1(c)
1826. Fabricius. K. Dansk. Vid. Selsk. Skr., ii, p. 57-
1841. Kiener. Cog. Viv. Marginella, p. 41, pl. 13, fig. 4 (zonata, preocc.).
1848. Krauss. Sidafrik. Moll., p. 126, pl. 6, fig. 22 (bilineata).
Radula large for the size of the animal: 1:5 x 0-3 mm. in shell 6 mm.
long; 60 rows, median tooth largest, flanked by 8-9 smaller teeth and 1-2
denticles on either side.
Off Tugela River (Natal-Zululand), 47 fathoms, one; off East London,
20 fathoms, 2; off Gove Rock (East London area), 22 fathoms, 2; off Keis-
kamma Point, 33 fathoms, one (S. Afr. Mus. P.F. coll.).
Marginella adela Thiele
1903. Von Martens. D. Tiefsee Exp., vii, p. 36 (not the fig. = multizonata = cylindrica).
1925. Thiele. loc. cit., p. 192, pl. 33 (21), fig. 23.
35, 16'S. 22° 26’ E., 155 metres (Sta. 104, corrected by Thiele, instead of
Sta. 114 von Martens).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 5
Off Cape Natal (Durban), 54 fathoms, 3; off Hood Point (East London
area), 5; 33 3 9- 27°57 E., 32 fathoms, 5; 34° 26’ S. 25° 42’ E., 125 fathoms,
several; off Cape St. Blaize, 125 fathoms, several (S. Afr. Mus. P.F. coll.).
The above P.F. specimens seem referable to Thiele’s species. None of
them show the four colour bands distinctly, though in two or three of them
_ there are traces of 1-2 bands.
Marginella musica Hinds
1844. Hinds. Proc. Zool. Soc. Lond., p. 73.
1844. id. <ool. Voy. Sulphur. Moll., p. 44, pl. 13, figs. 8, 9.
1848. Adams & Reeve. ool. Voy. Samarang., p. 28, pl. 7, figs. 4a-c, coloured (diadochus).
1886. Watson. Challenger Rep., xv, p. 265.
1903. Smith. Proc. Mal. Soc., v, p. 364 (diadochus).
1903. Von Martens. D. Tiefsee Exp., vii, p. 33 (diadochus).
1917. Tomlin. loc. cit., p. 263 (diadochus) and p. 283 (musica).
gee5- Uhiele. loc. cit., p. 194.
Cream, pale buff or grey, with narrow dark grey spiral lines varying in
number on last whorl from 5-13 (on base, i.e. length of aperture, varying from
4-10). Mature examples 15 to 25 mm. jong. Foot and mantle pale pink with
crimson radiating stripes (Adams & Reeve, fig. 4a).
B54 ©. 16 37 E., 150 fathoms (Watson); 33° 41 S. 18° E., 178 metres;
Sees re 21 E.., 916 metres; and 35° 16'S. 22°16’ E., 155 metres (von
Martens) ; St. Francis Bay (Thiele).
Off Cape Recife, 56-124 fathoms, Agulhas Bank and around Cape Point
to the Saldanha Bay area, 30-230 fathoms, one station in False Bay 14 fathoms
fear. Mus. P.F. coll.).
Living: Simon’s Bay (von Martens); False Bay (U.C.T.), 35° S. 20° 40’
E., 91 metres (s.s. Africana, per U.C.T.); Saldanha Bay area, 55 and 100
fathoms; off Cape Point 145 and 230 fathoms; and Agulhas Bank, 45 fathoms
(P2E. coll.).
Remarks. A characteristic and abundant species.
The largest specimen is 25 X 12°5 mm. (off Cape Point, 85 fathoms); the
smallest examples 5 X 3 mm., 4 X 2°5 mm., and 3 xX 2 mm.; the two former
have resp. 24 and 24 whorls, the latter has 2 whorls; all have 4 spiral lines on
last whorl. |
There are plump and slender forms: 11-5 X 7:5 mm. and 11°5 X 6 mm.,
22-5 X 12 mm. and 22-5 x 10°5 mm.; both these slender examples are from
the Saldanha Bay area, but not all the examples from that area are slender.
The type locality for musica is Cape Blanco, that for diadochus is Sunda
Strait. Watson said his Cape specimen had the more elongate form of diadochus,
but united this species with musica. Smith was inclined to agree. Thiele
regarded the Cape specimens as musica. ‘Tomlin listed them as separate species.
Dautzenberg (1910, 1912) does not record musica from West Africa, but
Knudsen does (1956. Atlantide Rep., 4, p. 91, pl. 2, fig. 17).
6 ANNALS OF THE SOUTH AFRICAN MUSEUM
The West African eveleighi Tomlin & Shackleford (1913. 7. Conch., xiv,
p. 11, pl. 1, figs. 5 and 6) strongly resembles specimens of this species with
numerous dark lines, especially Knudsen’s figure (1956. Atlantide Rep., 4, p. 84,
pl: 9, fie: 2)-
Turton’s fulvocincta may be a synonym, but is more likely to be a variation
of pzperata.
Marsinella mosaica Sow.
1846. Sowerby. Thes. Conch., i, p. 381, pl. 75, figs. 58, 59.
1892. id. Mar. Sh. S. Afr., p. 19 (var. langleyt).
A juvenile consisting of 2+ whorls, 4 x 2:5 mm., has one series of orange
dots above the shoulder, and 5 or 6 series below. It is relatively narrower than
the juveniles of bzcatenata.
Port Elizabeth (Sowerby); Port Alfred (Bartsch, Turton).
Off East London, 43 fathoms, one dead; off Nieca River (S. of East
London), 43-50 fathoms, one dead and one juv.; off Stalwart Pomt (N. of
Great Fish Point), 53 fathoms, one dead (S. Afr. Mus. P.F. coll.).
Marginella bicatenata Sow.
1914. Sowerby. Ann. Mag. Nat. Hist. (8), xiv, p. 477, pl. 19, fig. 7.
1916. Shackleford. Ann. S. Afr. Mus., xiii, p. 193, text-figs. 1, 2 (tomlini).
Creamy-white, body whorl with 2 rows of dark grey spots, one row around
the shoulder, and is visible also on the two preceding whorls of the spire, the
other row starting from the uppermost columella pleat.
Cape St. Blaize, N. x E.4E., 68 miles, 105 fathoms (Shackleford) ; Cape
St. Blaize, N. x E., 73 miles, 125 fathoms, one juv.; off Umkomaas River
(Natal), 40 fathoms, two juv. (S. Afr. Mus. P.F. coll.).
Remarks. The original locality was given as Goree (West Africa) with a
query; but possibly the specimen came from the P.F. collection sent to Sowerby.
The juveniles are 3-3 X 2°5 mm. with 2 whorls, 5 X 3°5 mm. with 24
whorls, and 5:5 X 3°75 mm. with 3 whorls; apex very blunt, rounded, angle
c. 100°; body whorl white with the distinctive 2 rows of spots, but faded. The
smaller juvenile is from the Natal locality.
A larger specimen, 22 X II mm., may be referable to this species. The
spire is higher, angle c. 60°, and the 4th and 5th whorls are irregularly and
shallowly fluted axially. Two rows of very faded spots can just be traced in the
same position as in the type of tomlini. P.F. coll., but no precise locality (S. Afr.
Mus. no. A 8776).
Marginella brocktoni Shackleford
1903. Von Martens. D. Tiefsee Exp., vii, p. 37 (chrysea, non Watson; not the fig. = perla).
1914. Shackleford. Ann. S. Afr. Mus., xiii, p. 98, 2 text-figs.
1925. Thiele. loc. cit., p. 193 (correction to von Martens).
y
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CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 7
34° 33'S. 18° 21’ E., 318 metres (von Martens). Cape Point, N. 50° E.,
180 fathoms, two (Shackleford).
The Pieter Faure obtained no other examples of this species.
Marginella augusta Thiele
1925. Thiele. loc. cit., p. 193, pl. 33 (21), fig. 28.
95° 11'S. 23° 2’ E., 500 metres (Thiele). One specimen, 17 X 6-5 mm.
(outer lip broken), without precise locality (S. Afr. Mus. P.F. coll.).
Remarks. Thiele’s figure shows 4 columella pleats, but in his description
he does not reckon the anterior fold of the columella as a pleat.
Very like brocktoni, but with 4 columella pleats.
Marginella neglecta Sow.
1846. Sowerby. Thes. Conch., i, p. 390, pl. 76, figs. 135, 136.
1852. Krauss. Arch. Naturg., i, p. 38 (reevei).
1853. Gaskoin. Ann. Mag. Nat. Hist. (2), xi, p. 359 (rufula).
1903. Von Martens. D. Tiefsee Exp., vii, pl. 3, fig. 3 (reevei) (not the specimens = clara).
1917. Tomlin. loc. cit., p. 283 (neglecta), p. 294 (reevet), p. 295 (rufula).
No specimens definitely referable to this species were obtained by the
Pieter Faure.
Marginella clara ‘Thiele
1903. Von Martens. D. Tiefsee Exp., vii, p. 35 (not the fig. = reevei = neglecta).
moet Einele. loc. cit., p. 192, pl. 33 (21), figs. 21, 22.
35, 16'S. 22° 26’ E., 155 metres (von Martens, Thiele).
Very like neglecta, or possibly a not fully mature atractus.
Marginella bensoni Rve.
1865. Reeve. Conch. Icon., pl. 27, fig. 158.
1904. Smith. 7. Malac., xi, p. 32, pl. 2, fig. 20 (dulcis).
? 1925. Thiele. loc. cit., p. 195, pl. 33 (21), figs. 35, 36 (laetztia).
A faint spiral band below the suture and another on lower part of base
(Thiele). Thiele said his species was near adela, but smaller. It certainly seems
near bensont.
By 10'S. 22° 26’ E., 155 metres (Thiele).
P.F. specimens from the following localities seem referable to bensonz;
they are smaller than Thiele’s specimens.
Off Umkomaas River (Natal), 40 fathoms; off Cape Morgan, 47 fathoms;
34° 55'S. 25°55 E., 67 fathoms; False Bay, 22 fathoms (S. Afr. Mus. P.F.
coll.).
8 ANNALS OF THE SOUTH AFRICAN MUSEUM
Marginella differens Smith
1892. Sowerby. Mar. Sh. S. Afr., p. 20 (bulbosa, non Reeve).
1904, Smith. 7: Malac.. xt p. 32, pi--2, fig..19:
1916. Shackleford. Ann. S. Afr. Mus., xiii, p. 194, text-figs. 3, 4 (taylort, non Olsson).
1919. Tomlin. Proc. Mal. Soc., xiii, p. 65 (barnardi, nom. nov. for taylori, preocc.).
The type and cotype of taylort in 8. Afr. Mus. do not seem distinguishable
from other specimens identified by Shackleford as differens.
differens forma eugenes n.
Shape of differens, but larger and slightly less obese. 4 whorls. Columella
pleats 4 with traces of 1-4 additional pleats posteriorly. Outer lip not reaching
suture above, scarcely shouldered, not varicoid, internally with 18-20 well-
developed denticulations or plicae, slightly inflected at upper end, aperture
indented anteriorly. 7-5 x 4:8 mm. (Type), another 8 x 5 mm. Uniform
pale buff.
Off Cape Natal (Durban), 85 fathoms, one; off Umkomaas River (Natal),
40 fathoms, one; 34° 27'S. 25° 42’ E., 256 fathoms, 5; 34° 26 sympa
124 fathoms, 6; Gericke Point (Knysna area), 42 fathoms, one; off Cape St.
Blaize, 37 fathoms, 2; the same, 125 fathoms, 11; off Cape Infanta, 46 fathoms,
one (5) Air Mins: weaker colle
Remarks. ‘The 8 xX 5 mm. specimen from off Gericke Point was seen by
Tomlin and Shackleford, and considered to be a n. sp. but ‘too poor’ for
description. Actually it seems in quite fair condition, though the surface is not
glossy. Therefore the glossy, but slightly smaller, specimen from off Umkomaas
River is taken as the type (S. Afr. Mus. no. A8786) of this large form.
Only further research will show whether a name is really necessary; the
difference in size between this form and normal differens (5-8 mm.) is no greater
than that found in musica (15-25 mm.).
Marginella burnupi Sow.
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 10, pl. 6, fig. 35.
1929. Dautzenberg. Faune Col. Franc., 111, 4, p. 381.
1932. Turton. Mar. Sh. Port Alfred, p. 42.
Not Thiele. 1925. loc. cit., p. 192, pl. 33 (21), figs. 24, 25.
Conical, widest slightly above middle, narrowing anteriorly, spire flat.
Columella pleats 5 (Sowerby), 6-8 in larger specimen. Outer lip incrassate
(when mature), internally plicate. 4 <x 2 mm. (Sowerby); 5 X 3 mm.
Port Elizabeth (Sowerby); Port Alfred (Turton).
Algoa Bay, 67 fathoms, one, typical; off Glendower Beacon (Port Alfred
area), 66 fathoms, one, with low spire (S. Afr. Mus. P.F. coll.).
Distribution. Madagascar (Dautzenberg fide Bavay).
Remarks. Of 4 worn specimens from Port Alfred (coll. Turton) one
(immature) has a flat spire, the others have low spires, definite but not so high
as in Thiele’s figure.
;
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;
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CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 9
Thiele had difficulty with burnupi and dulcis (= bensoni) owing to variabi-
lity, and suggested that the plump and slender forms might possibly be the
two sexes. Both his figures show rather high spires, quite unlike Sowerby’s
figure and the typical specimen recorded above; moreover, fig. 24 shows a
denticulate lip (which is correct for burnupi), but fig. 25 shows a non-denticulate
iip. Thiele’s specimens came from Algoa Bay, Agulhas Bank, and St. Francis
Bay; also from Great Fish Bay (Angola). I doubt whether any of these were
really burnupi; they should be re-examined, especially the Angolan specimen.
M. almo Bartsch 1915 also has a low spire, but is ovoid in shape with the
greatest width in the middle.
Marginella fallax Smith
1903. Smith. Proc. Mal. Soc., v, p. 365, pl. 15, fig. 20.
Off East London, 32 fathoms, two; False Bay, 20 fathoms, 3 (S. Afr. Mus.
EF, coll.).
A worn specimen in 8. Afr. Museum is said to have come from ‘Natal’,
but the record is unreliable.
Marginella keen Marrat
1871. Marrat. Ann. Mag. Nat. Hist. (4), vil, p. 141, pl. xi, fig. 13.
1925. Thiele. loc. cit., p. 194, pl. 33 (21), fig. 30 (agulhasensis).
Bae 22 96° E., 155 metres (Thiele).
Cape Point N. 16° E., 10 miles, 85 fathoms, 2; off Glendower Beacon
(Port Alfred area), 100 fathoms, one (S. Afr. Mus. P.F. coll.).
Remarks. Not having seen Marrat’s original description, I accept Tomlin
and Shackleford’s identification of the three P.F. specimens. They are clearly
the same as Thiele’s agulhasensis.
The 2 Cape Point specimens (13 < 7 mm.) have 3 columella pleats, one
of them with an additional indistinct one as shown in Thiele’s figure; the
Glendower Beacon specimen has 4 well-defined pleats. The latter is somewhat
worn, and hence proportionately broader (11 X 7 mm.).
Marginella shepstonensis Smith
1906. Smith. Ann. Natal Mus., i, p. 31, pl. 7, fig. 5.
Wavy or zigzag brown axial lines, thickened in two or three places so as
to form 2 or 3 spiral bands of crescentic marks, one in the middle and one
towards each end of the body whorl.
Off Tugela River (Natal-Zululand), 47 fathoms, one; off Illovo River
(Natal), 27-30 fathoms, 2; off Umkomaas River (Natal) 40 fathoms, one
(S. Afr. Mus. P.F. coll.).
10 ANNALS OF THE SOUTH AFRICAN MUSEUM
Marsginella kerochuta Shackleford
1903. Von Martens. D. Tiefsee Exp., vii, p. 34 (not the fig. = zeyhert).
1914. Shackleford. Ann. S. Afr. Mus., xili, p. 97, 2 text-figs.
1925. Thiele. loc. cit., p. 191 (correction to von Martens).
Biconical, outer lip when mature forming a strongly projecting shoulder,
spire (including shoulder) subtending an angle of 70° (type) to go°. Mature
9 X 6b te 13 x 7 mm. (Phicle? lone. 6-11 mm_)-
34° 39'S. 18° 21’ E., 318 metres (von Martens); 35 16 S27 22 -seeeee
155 metres (Thiele).
Off Cape Point, 135 fathoms (Shackleford).
36° 40’ S. 21° 26’ E., 200 fathoms, one fresh; off Cape St Blaizemmeg
fathoms, 6 worn; Brown’s Bank (approx. 364° S. 21° E.), 80-100 fathoms, one
fresh) amimature (o. Afr. Mus, P.F. coll):
Remarks. Type and cotype in 8. Afr. Mus. The type has the highest spire
with angle 70°, the cotype with angle 80°; the specimen from southern end of
Agulhas Bank with angle 80°, and the worn specimens with angles 85°—g0°.
Closely related to zeyheri, but larger and from deeper water.
Mareginella zeyhert Krauss
1852. Krauss. Arch. Naturg., i, p. 38.
1892. Sowerby. Mar. Sh. S. Afr., p. 20 (metcalfei, non Angas).
1903. Von Martens. D. Tiefsee Exp., vii, pl. 3, fig. 4 (Krauss type) (not the description =
kerochuta).
1904. Smith. 7. Malac., xi, p. 31, pl. 2, fig. 18 (pura).
1917. Tomlin. loc. cit., p. 306, and p. 292 (pura).
1925. Thiele. loc. cit., p. 191 (correction to von Martens).
1925. id. ibid., p. 195, pl. 33 (21), figs. 33, 34 (aurelia).
Biconical, outer lip forming a strongly projecting shoulder, spire angle 70°.
Mature: 5 X 3 to 7°5 X 4-4°5 mm.
Port Elizabeth (Sowerby); Port Alfred (Smith, Bartsch, Tomlin).
35° 16'S. 22° 26’ E., 155 metres; 34° 8’ S. 24° 50’ E., 80 metresm aimee:
Off Umkomaas River (Natal), 40 fathoms, 2; off Cape Morgan, 47
fathoms, 2; off Cove Rock (East London area), 22 fathoms, 5; off East London,
20 fathoms, 6 (S. Afr. Mus. P.F. coll.).
Remarks. Specimens in which the outer lip is worn smooth have the
appearance of belonging to the non-denticulate section of the genus.
Marginella atractus Tomlin
1903. Sowerby. Mar. Invest. S. Afr., li, p. 227 (fusiformis, non Hinds).
1918. Tomlin. 7. Conch., xv, p. 306, pl. 10, fig. 6.
1925. Thiele. loc. cit., p. 195, pl. 33 (21), figs. 31, 32 (julia).
1925. id. ibid., p. 195, pl. 34 (22), fig. 1 (meta).
Fusiform, outer lip when mature forming a well-marked shoulder, spire
angle 50°. 3 X 1°5 mm. (2$ whorls); 4 x 1°75 mm. (3 whorls); mature:
5°5 X 2°5 mm. to 8 X 3°75 mm.
wt cP ee
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA II
Nanquas Peak (eastern end of Algoa Bay), 49 fathoms (Sowerby); Port
Elizabeth (Smith); Port Alfred (Tomlin, Turton); Still Bay (S. Afr. Mus.
Muir coll.).
wees: 19° 97 E., \So metres; 95° 16'S; 22°26. E:, 155: metres;
epee 5. 21 3 F., 102 metres; and 35° 26'S. 20° 56’ E.sdepth ? (Thiele).
Off East London and Cove Rock, 22-32 fathoms; off Keiskamma River
and Great Fish Point; Algoa Bay 22-30 fathoms; off Cape Recife, 125 and
256 fathoms; off Cape St. Blaize, 125 fathoms (S. Afr. Mus. P.F. coll.).
Remarks. Judging by the numbers of specimens retrieved from the P.F.
bottom-samples, this species seems concentrated chiefly in the East London to
Algoa Bay area.
Over 100 examples have been examined. Comparatively few specimens
show a definite lip denticulation; often it is traceable only in an oblique light;
much depends on the condition of the specimen.
The proportions vary slightly, though the present material does not
_ altogether support Turton’s statement that the broader zeyheri and narrower
atractus ‘grade together’ (beach specimens may do so!); I have found little
difficulty in separating them and therefore maintain the two species. On the
other hand the slight differences relied upon by Thiele for his species do not
seem to justify additional species.
Marginella perminima Sow.
Fig. 1(d)
1894. Sowerby. 7. Conch., vii, p. 370.
1897. id. Append. Mar. Sh. S. Afr., p. 9, pl. 6, fig. 36.
1932. Turton. loc. cit., p. 43 (referred to under neptunz).
Columella pleats 4 plus 7-8 tiny denticles.
Port Elizabeth (Turton).
Off Cape Natal (Durban), 54 fathoms, one; off Umkomaas River (Natal),
40 fathoms, one; off Sandy Point (N. of Cape Morgan), 51 fathoms, 2; off
Cove Rock (East London area), 22 fathoms, 2; 33° 3’ S. 27° 27’ E., 32 fathoms,
one; 34° 27'S. 25° 42’ E., 56 fathoms, one; off Cape St. Blaize, 125 fathoms,
ene (o. Afr. Mus. P.F. coll.):
Remarks. ‘The original description is not available to me. Turton says the
species has only 3 columella pleats. In some of the present specimens only 3
are Clearly visible, the fourth and the additional denticles being seen only in
the best specimens.
Fam. CANCELLARUDAE
1903. Thiele D. Tiefsee Exp., vii, p. 171 (radula).
Ig11. Schepman. Siboga Exp. monogr., xlix, 1, p. 265 (radula) (line 18 for ‘nearly’ read
‘neatly’).
1955. Adam & Knudsen. Bull. Inst. Roy. Sc. Nat. Belge, xxxi, no. 61, p. 18 (radula).
1958. Barnard. 7. Conch., xxiv, p. 243 (radula).
12 ANNALS OF THE SOUTH AFRICAN MUSEUM
Eight species have been known hitherto from South African waters. In
the present paper one new species and one new record are added.
C. dalli Bartsch 1915 is known from a single specimen recorded as coming
from the Cape of Good Hope. C. plebeja Thiele 1925 is known from two shells
from the Agulhas Bank. The Challenger species imbricata Watson 1882 was
rediscovered by the Pieter Faure, which also obtained the new species, but no
examples of plebeja.
The most interesting discovery, however, was made by the Fisheries
Survey vessel Africana off Liideritzbucht, viz. a species which seems referable
to the Italian Pliocene-Miocene lyrata. Specimens identified with this fossil
species have already been recorded from off the Cape Verde Islands and the
coast of northern Angola, in the latter locality living. The extension southward
of the range of this species, indeed its presence in three localities off the west
coast of Africa, raises very interesting zoogeographical questions.
In three species I have been able to make some observations on the
remarkable radula of the genus Cancellaria. Unfortunately the condition of the
material left no opportunity of investigating the myology.
The radula consists of a large number (at least 100) of very long slender
teeth attached in single file to a basal membrane. The teeth are oriented in
two groups: those attached to the shorter anterior portion of the membrane
project forwards, the more numerous teeth on the posterior portion project
backwards, and appear to be replacers.
The radula is enclosed in a double sheath (Barnard, loc. cit., fig.): a
smaller inner one rather like the carapace of a bivalve Crustacean; the outer
larger one has a pointed tubular anterior projection with a small apical opening.
I have suggested that possibly this buccal apparatus operates by pushing the
anterior teeth through the tubular opening of the outer sheath, which would
hold the teeth firmly like the hairs in the collar of a paint-brush, and thus
using them to sweep particles of food into the mouth. But the myology should
be studied, and for this purpose fresh material is necessary.
Cancellaria imbricata Watson
Fig. 2
1882. Watson. 7. Linn. Soc. Lond., xvi, p. 325.
1886. id. Challenger Rep., xv, p. 274, pl. 18, fig. 10.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 230.
1958. Barnard. loc. cit., p. 244.
Protoconch (corroded) smooth, diam. 2 mm. Postnatal whorls 4; 1st
with 3 spiral lirae, 2nd and following whorls with 4, crossed on 1st and and,
and sometimes part of 3rd whorl by feeble axial ribs, intersections slightly
tuberculose. Growth-lines closely imbricate, retractively concave between each
pair of spiral lirae. 10-12 additional lirae on base. 29 X 17 mm., a juv.
13 X 8mm.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 13
Radula with at least 100 extremely long slender filiform teeth set closely
together in two divergent series on a narrow basal membrane; apex of each
tooth truncate, obscurely denticulate, one of the margins near apex also
obscurely denticulate. Length of basal membrane c. 1 mm., of each tooth
c. 2mm.
Shell dull white; animal (as preserved) greenish.
35° 4'S. 18°.37’ E., 150 fathoms, green sand (Watson). Off Cape Point,
135-190 fathoms, green sand with black specks (S. Afr. Mus. P.F. coll.).
Se a ee
Fic. 2.
Cancellaria imbricata Watson, face (slightly oblique) and lateral views of radula plates on basal
membrane, with apex of one plate further enlarged.
Remarks. The Challenger obtained one specimen; the Pieter Faure 14 speci-
mens (including the one recorded by Sowerby); three were taken alive.
The apex of all the shells, including the juvenile of 13 mm., is more or less
corroded.
Although Watson said the aspect of the shell suggested an Admete, the
presence of a radula shows that he placed the species in the correct genus.
Cancellaria bifasciata Desh.
1859. Chenu. Man. Conchyl., i, p. 277, fig. 1845.
Umbilicus narrowly open. Aperture longer than spire. Whorls rounded,
suture deep, but visible laterally. Protoconch 2 whorls, smooth. Postnatal
whorls 34; at first spirally lirate, then growth-lines becoming stronger and
forming numerous axial riblets on 2nd and first part of 3rd whorl, producing
a cancellate sculpture; but on body whorl becoming again subordinate to the
spiral lirae; the latter c. 11 on body whorl, with a fine intermediary between
each of the upper 3 or 4 pairs of main lirae; ¢. 16 additional lirae on base,
very regularly arranged. Columella with 3 pleats. Aperture oval, no canal.
24 X 14 mm.
14 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pale buff with faint orange patches above and below a pale band slightly
below middle of whorl.
Off Cape Natal (Durban), 47 fathoms, one complete; and 54 fathoms,
three broken body whorls (S. Afr. Mus. P.F. coll.).
Remarks. The complete specimen described above was identified by
Tomlin as bifasciata var., with the suggestion that further examples might show
it to be a distinct species. As far as can be judged, it is exactly like Chenu’s
figure. It also corresponds with a figure of elegans Sow. (Reeve. Conch. Icon.,
x, pl. 16, fig. 75) in Gravely (1942. Bull. Madras Govt. Mus., n. s., V, 2, p. 68
(in key), fig. 12 2).
The broken body whorls agree in sculpture with the above described
specimen.
Cancellaria euetrios n. sp.
Fig. 3
Umbilicus narrowly open. Aperture a little longer than spire. Whorls
convex, without shoulder, suture deep, not quite visible in lateral view.
Protoconch 14 whorls, alt. 0-5 mm., diam 0-75 mm., smooth. Postnatal
whorls 2, junction with protoconch not clearly
defined but marked by 4-5 indistinct and
incomplete axial ribs; 28 ribs on Ist, 30 on 2nd
whorl ; crossed. by spiral lirae 9-10 on Ist, 11 on
2nd whorl, 11-12 additional lirae on base.
Columella with 3 indistinct pleats.
4°5 X 2°5 mm.
White beneath pale buff periostracum,
protoconch white.
34° 26'S. 25° 49’ ER. (off Capemiecie.
124 fathoms, one (S. Afr. Mus. no. A8747.
PEs colle)p
/ Remarks. The size of the protoconch
i indicates a small species, but this specimen is
! regi probably not fully grown. The cancellate
; le sculpture is very regular (‘well-woven’) though
Fic. 3. perhaps not more so than in some other species;
Cancellaria euetrios n. sp. the ribs are more prominent than the lirae.
Cancellaria producta Sow.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 220, pl. 4, fig. 5.
Off Umhloti River (Natal), 40 fathoms, 2 specimens (Sowerby). Same
locality, 2 and one broken; off Umhlanga River (Natal), 22-26 fathoms, one
(S. Afr. Mus. P.F. coll.).
a
SS ee
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA BS
Remarks. One of the two type specimens, agreeing with Sowerby’s measure-
ments, is in S. Afr. Museum; the other ‘type’ was probably in coll. Sowerby
(? now Brit. Mus.).
Another specimen is broken and bored by Cliona, only the last 24 whorls
remaining; it was a larger shell than the type: width 8 mm., with presumably
long. c. 19 mm.
Cancellaria lamellosa Hinds
1843. Hinds. Proc. Zool. Soc. Lond., p. 49.
1844. id. Kool. Sulphur. Moll., p. 43, pl. 12, figs. 15, 16.
Ig1i. Schepman. Siboga Exp. monogr., xlix, p. 264.
1958. Barnard. loc. cit., p. 244.
Pale buff, a pinky-brown band at shoulder and another about in middle
of body whorl, more distinctly visible inside the aperture. One of Burnup’s
Durban specimens is orange-brown, the bands visible only inside the aperture.
The Delagoa Bay shell is very pale pink, with a few brown specks on the crest
of each axial rib. Up to 18 x 13 mm. Animal (as preserved) flesh tint.
Radula similar to that of imbricata.
Agulhas Bank (Hinds). Off Umvoti, Umhloti, and Umhlanga Rivers
(Natal), 22-40 fathoms (S. Afr. Mus. P.F. coll.). Durban (S. Afr. Mus. coll.
Burnup).
Living: Delagoa Bay (U.W.).
Distribution. Ceylon, Straits of Malacca, East Indies.
Remarks. The largest specimen (as above) is from Delagoa Bay. The
extent to which the umbilicus is covered by the columella callus is variable;
and also depends on the angle from which the shell is viewed. This explains
the words ‘umbilico magno’ in Hinds’s description.
In Gravely (1942. Bull. Madras Govt. Mus., n. s., V, 2, p. 68 (in key))
lamellosa is regarded as one of the varieties of crispa Sow.
Cancellaria foveolata Sow.
1848. Sowerby. Thes. Conch., i, pl. 93, figs. 30, 31.
1932. Turton. Mar. Sh. Port Alfred, p. 30, pl. 6, no. 224.
Pale buff or amber to chestnut-brown, sometimes irregularly infuscate or
with a brown band in middle of body whorl, brown spiral bands often visible
inside aperture, protoconch white. 24 x 14 mm.
Natal to Jeffreys Bay (S. Afr. Mus.).
Remarks. Not taken by the Preter Faure, and known only from beach
material.
Cancellaria semidisjuncta Sow.
1848. Sowerby. Proc. Zool. Soc. Lond., p. 137; and Thes. Conch., i, pl. 95, figs. 62, 63.
18 X 14 mm.; width of a larger specimen 18 mm., apex broken but full
length probably c. 25 mm.
16 ANNALS OF THE SOUTH AFRICAN MUSEUM
East London to Jeffreys Bay (S. Afr. Mus.).
Remarks. Not taken by the Pieter Faure. A specimen in 8. Afr. Mus., said
to have come from Tanganyika, is very worn and has the coarse cancellate
appearance of worn South African examples. Sowerby in his original descrip-
tion gave Philippine Islands as the locality, but Bartsch 1915 doubted this
locality. |
Cancellaria cf. lyrata (Brocchi)
Fig. 4
1814. Brocchi. Conch. foss. subapenn., ii, p. 311, pl. 3, fig. 6 (Voluta 1.) (quoted from D. & F.).
1820. Borson. Mem. R. Ac. Sc. Torino, xxv, p. 210 (quoted from Sherborn. Index Anim.).
1894. Sacco. Moll. Terz., part 16, p. 59, pl. 3, figs. 57-65 (Sveltza 1.) (quoted from D. & F.).
1906. Dautzenberg & Fischer. Res. Sci. Camp. Monaco, fasc. 32, p. 17, pl. 1, figs. 11-13
(Recent), figs. 14-16 (fossil).
1955. Adam & Knudsen. Bull. Inst. Roy. Sc. nat. Belge, xxxi, no. 61, p. 16, pl. 2, figs. 6, 7,
and text-fig.
1958. Barnard. 7. Conch., xxiv, p. 243 (radula).
Specimen A.
Aperture very little longer than spire. Protoconch tip broken, remainder
slightly corroded, diam. 1 mm. Postnatal whorls 5, profile of 2nd and grd
slightly angular, of 4th and 5th sharply keeled in middle; 1st whorl corroded,
2nd partly corroded but with indications of 9 or 10 (possibly 11) axial ribs,
grd with 12, 4th with 11, 5th with 9 ribs, from suture to suture and extending
across base; ribs narrow, sharp, with acute points at intersections with the
peripheral keel; crossed by spiral lirae, corroded on early whorls, at least a
dozen fine and equal-sized above periphery on 4th and 5th whorls, below
periphery 2 stronger main lirae and 2 weaker, with finer intermediaries; 6—7
additional lirae with intermediaries on base (total about 15); the main lirae
form little prickles at intersections with the ribs. Columella with 3 pleats,
columellar glaze not fully developed, thin and not obscuring the ribs and lirae
on base. Umbilicus closed. Aperture wide, outer lip thin, not constricted
below; canal short and wide. 35 x 24 mm. (incl. peripheral points, 19 mm.
excl. these points).
Greyish-white, aperture yellowish-fawn.
Locality? (see Remarks, znfra) (S. Afr. Mus. ex coll. P. Ross-Frames).
Specimen B.
Aperture a little longer than spire. Apex corroded. Remaining postnatal
whorls 4, profile angular, last whorl with peripheral keel. Early whorls
corroded, axial ribs 11 on each of the upper two whorls, 9 on last whorl, more
or less tubercular at shoulder, but corroded; crossed on last whorl by spiral lirae
(not traceable on preceding whorls), at least a dozen fine and equal-sized lirae
above shoulder, and about a dozen below, 3 or 4 slightly stronger than the
others, about 15 (partly corroded) additional lirae on base, varying in size.
1
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA by |
Columella with 3 pleats, columellar glaze extensive, rather thick, the axial ribs
and some of the lirae only indicated. Umbilicus closed. Aperture wide, outer
lip thin, not constricted below; canal short and wide. 43 X 24 mm.
Dirty brownish-grey, aperture yellowish-brown, glaze whitish.
Radula and sheath: see Barnard, loc. cit., and remarks under genus.
* 26° 26'S. 14° 39' E., 174 metres (off Liideritzbucht, South West Africa)
(s.s. Africana, 1948, AFR. 1260 B. per U.C.T.).
Distribution (recent). Off Cape Verde Islands, 628 metres (Dautzenberg
& Fischer); 5° 46’-6° 29'S. 11° 32’-11° 38’ E., 145-230 metres (Adam &
Knudsen).
Fic. 4.
Cancellaria cf. lyrata (Brocchi). Specimen A on left, B on right; with diagrammatic apical
views of last whorl.
18 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. Unfortunately the provenance of the less corroded Ross-Frames
specimen (A) is unknown. The late P. Ross-Frames collected many shells
while serving in the military campaign in South West Africa in 1914-15 (cf.
Burnup. 1923. Ann. Natal Mus., V, p. 2), and in view of the larger living
example (B) having been dredged off Liideritzbucht, it is a reasonable assump-
tion that he obtained his specimen, directly or through a friend, from somewhere
in the same area.
The two specimens are obviously conspecific. The Ross-Frames specimen
is the less corroded, and is much fresher-looking. It is slightly smaller, and the
columellar glaze has not reached its full extent or thickness. It shows, however,
that the remaining whorls on the rather badly corroded Africana specimen are
actually the 1st to 5th postnatal whorls, not a corroded protoconch plus 4
whorls.
There is a striking resemblance between these shells, especially the Africana
one, and the shell dredged off the Cape Verde Islands, which Dautzenberg &
Fischer identified with the Italian Pliocene-Miocene lyrata. There is an even
greater, in fact an exact resemblance to the specimens described and figured
by Adam & Knudsen from off the northern part of the Angolan coast, and
likewise referred to the same fossil species.
The fewer and more widely spaced axial ribs on the last whorl in both
specimens A and B should be borne in mind in deciding the status, specific or
variational, of the South West African form. The fossil form is known to be
variable.
Cancellaria lyrata Ad. & Rve. (1850. Zool. Samarang. Moll., p. 42) is probably
not the same as lyrata (Brocchi), and if so requires renaming.
Fam. VOLUTIDAE
1901. Smith. Proc. Mal. Soc., iv, p. 231 (synopsis of S. African species).
1922. Cooke. ibid., xv, p. 6 (radulae).
Gen. VoLuTa Linn.
Operculum present or absent. Columella with pleats. Radula with
tricuspid central tooth only.
Of the five endemic South African species, the radulae af three are known,
and one is definitely known to possess an operculum.
Spire high, pointed. With axial ribs.
Columella callus black’ 2). 20. 0 Ue ee
Callus not black.
Pleats 3 plusg obscureones .».+ 3) (aia eens i
PHC AtS a ee a) nae
Spire low, blunt. No axial ribs.
Pleats). a ee arte da MOT ee Ca
Pleats 4 en else gy IN, UE ee ee. ara Se
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 19
V. mitraeformis, V. scapha, and Melo armata were not admitted to the fauna-
list by Smith (1901). I would also exclude V. festiva; if the young specimen
recorded by Sowerby (1897) was really found on the Natal coast, it was far
more probably an africana or a ponsonbyt.
Voluta africana Rve.
1856. Reeve. Proc. Zool. Soc. Lond., p. 2, pl. 33, figs, 3, 4.
1903. Von Martens. D. Tiefsee Exp., vii, p. 31.
1913. Bullen Newton. Rec. Albany Mus., ii, p. 342, pl. 24, figs, 1,2.
1922. Cooke. loc. cit., pp. 7, 10, fig. 3 (radula).
1932. Turton. Mar. Sh. Port Alfred, p. 44 and vars. beckeri, ponderosa, pl. 9, no. 327 (juv.),
no. 328, and pl. 10, no. 329.
1933. id. 7. Conch., xix, p. 370 (rietensis, nom. nov. pro ponderosa, preocc.).
Spire high, pointed. Aperture 13-13 times spire. Protoconch 24 whorls,
diam. 4-4-5 mm. (3:5 mm. on an old worn specimen). Postnatal whorls 3.
Axial ribs strongly tuberculate at shoulder (but usually abraded), extending
_ below to base in juveniles (up to 16 mm.) but in large specimens becoming
obsolete (except in vars. beckert and rietensis), 16-18 on 1st whorl, 9-11-13
(sometimes 14) on last whorl; spiral striae over whole whorl, but usually
obsolete on body whorl except in fresh specimens. Columella pleats 5, the
upper 2 obscure. Outer lip in adult slightly thickened and ascending towards
shoulder of preceding whorl. A well-marked columella callus. 67 x 39 mm.
An operculum ‘presumably belonging’ to a specimen of this species was
described by Smith.
Pinky-brown, more or less speckled, speckles usually aggregated to form
two more or less solid spiral bands, sometimes with large brown blotches; dark
brown spiral lines may appear towards outer lip, but usually they are only
visible on the thickened lip where they occur in pairs, threes, or fours; colu-
mella callus dark chestnut-brown (Smith: ‘coal-black’). Juveniles with 4-5
narrow spiral bands on Ist whorl (or first and a half), continuous or broken
into a series of spots (cf. Turton’s fig. 327).
Radula with c. 54 rows (Cooke).
Fossil: Mio-Pliocene: Redhouse near Port Elizabeth (Newton).
East Africa (Reeve); South-east Africa (Sowerby); Port Elizabeth
(Sowerby); Port Alfred (Bartsch, Turton); Port St. Johns and Pondoland
(Smith); off Durban, 40 fathoms (Smith); 35° 16’ S. 22° 26’ E., 155 metres,
fragmentary lower half, 38 mm. long (von Martens). Natal, from fish stomachs,
with var. beckert; Port Alfred, juveniles and var. rietensis (S. Afr. Mus.).
Not taken by the Preter Faure.
var. beckeri (Shackleford ined.) Turton. A narrow form with usually more
numerous ribs (14) which extend to base, sometimes 2 ribs concurrent into one
tubercle (e.g. 14 ribs, 10 tubercles), tubercles less prominent. Occurs in Natal
along with the plump form.
var. rietensis Turton. A thicker-shelled, heavier form, with ribs extending
to base. Maybe these shells are merely aged individuals. 68 x 45 mm. (badly
20 ANNALS OF THE SOUTH AFRICAN MUSEUM
worn), 73 X 46 (apex and canal worn) (S. Afr. Mus.). Recorded only from
Port Alfred.
Remarks. If the operculum described by Smith really belonged to this
species, the species cannot be placed in the subgen. Alcithoe.
Both plump and slender forms occur: 38 X 21, 51 X27) uhemeueg,
64 X 35, 64 x 37, 65 X 35, 67 X 39 mm.
S. Afr. Mus. has protoconchs and juveniles up to 16 mm., but no speci-
mens between 16 and 38 mm. long.
Specimens with thickened outer lip (with dark lines) 38 and 42 mm. long.
Voluta ponsonbyt Smith
1901. Smith. loc. cit., p. 231, text-fig.
1922. Cooke. loc. cit., pp. 7, 10, fig. 4 (radula).
Spire high, pointed. Aperture 1$-1? times spire. Protoconch 24 (3)
whorls, diam. 2-2-5 mm. Postnatal whorls 4. Axial ribs sharply pointed at the
shoulder producing a coronate appearance, extending only a short distance
towards base on body whorl, 17-20 (22) on 1st whorl, decreasing to 10-14 on
last whorl; on 1st whorl spiral striae on upper part and in the intervals between
the ribs (if unworn extending across the ribs), on later whorls visible only on
upper part (shoulder to suture). Columella with 6 pleats, the upper 3 obscure.
Outer lip in adult thickened and ascending towards shoulder of previous
whorl. A well-marked columella callus. 83 x 39 mm., others 74 X 36,
78 X 42 mm.
Operculum?
Salmon coloured, with white spiral bands, 7 between shoulder and base
on body whorl, crossed by darker lines, two of the broader intervening salmon
bands with brighter salmon or orange-brown patches, sometimes somewhat
irregular; a dark spot on front of each tubercle; columella and interior of outer
lip pinkish, callus white. |
Radula with ¢. 53 rows (Cooke).
Off Durban, 40 fathoms (Smith). Natal coast, from fish stomachs. (S. Afr.
Mus.). Not taken by the Pieter Faure.
Remarks. Smith gave the distinctive differences between this species and
festiva, size being one of them; but he omitted the sizes of the respective proto-
conchs. Comparisons as regards size are risky. Smith did not give the size of
festiva, but said ponsonby: was the smaller species. His specimen with thickened
outer lip, therefore presumably mature, was 57 mm. long (there is an exactly
similar sized one in 8. Afr. Mus., also with thickened lip); but this is greatly
exceeded by several specimens in S. Afr. Mus., viz. (with thickened lip and
more or less developed callus) 54 (2 specimens), 57, 59, 60, 64, 65 (2 specimens),
74, 78 and 83 mm.
SE a ee ee
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 21
There is a tendency in some specimens for the tubercles to become obsolete
on the later part of last whorl; in one specimen the tubercles cease abruptly
(only g instead of 11 or 12), and the profile of this part of the whorl is evenly
curved with scarcely any shoulder.
The dimensions of the 3 largest specimens show that there are plump and
_ slender individuals.
Has been taken only from fish stomachs.
Voluta queketti Smith
Fig. 5
1g01. Smith. loc. cit., p. 234, text-fig.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 226.
1922. Cooke. loc. cit., pp. 7, 10, fig. 6 (radula).
Spire pointed, aperture 13-1? times spire. Protoconch 24 whorls, diam.
I°75-2 mm. Postnatal whorls 5. Axial ribs 19-21 on 1st whorl, decreasing to
11-14 on last whorl, extending from suture to suture and 2 towards base on
body whorl, sharp, especially in juvenile, at top forming projecting points
(especially sharp in juvenile), subcoronate and concealing the sunken suture,
but rounded off on the last whorl leaving the suture exposed; numerous close-
set spiral striae over whole whorl, becoming more widely separated on base,
especially well marked at the tops of the ribs in juvenile. Columella pleats 6
(or 7), the upper one or two feeble, in the largest specimen 6 distinct pleats.
Outer lip in largest specimen somewhat exsert, sharp edged. 51 X 22 mm.
Operculum narrow-oval, 11 X 4:25 mm. in aperture 30 mm. of 51 mm.
shell, nucleus a little distance from apex and nearer the outer than the inner
margin.
Fic. 5.
Voluta queketti Smith. Operculum of 51 mm. shell. Second and upper part of 3rd whorl of
16 mm. juvenile.
22 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pale yellowish flesh-tint (Smith) with bright red blotches on upper ends of
ribs, an irregular interrupted red band above middle of body whorl and some
spots on lower part. Smith’s description applies to the present specimens. The
largest one, taken alive, has the internal margin of outer lip pinkish, columella
white; operculum horn colour.
Radula with ¢. 41 rows (Cooke).
Off Durban, 40 fathoms (Smith); off Cape Natal (Durban), 27 [sic = 55]
fathoms; off O’Neil Peak (Zululand), 90 fathoms; off Umhloti River (Natal),
27 [sic = 54] fathoms (Sowerby).
Off Umhloti River (Natal), 40 fathoms, one living; off Cape Natal, 54—
55 fathoms, 7 dead (one 36 mm. long, 4 juv. and 4 broken); off O’Neil Peak,
55 fathoms, one dead (34 mm. long) and one broken (S. Afr. Mus. P.F. coll.).
Remarks. Plump and slender forms occur: 37 < 17 (Smith’s type),
36 X 19 and 34 X 15 mm. (S. Afr. Mus.). S. Afr. Mus. has juveniles 14 mm.
(protoconch incomplete) to 20 mm. long.
Smith had one specimen; Sowerby returned 7 P.F. specimens, now in S. Afr.
Mus., but apparently retained the O’Neil Peak specimen from go fathoms
(unless the depth is wrongly recorded, as in his other two records). No record
is available of how many specimens were sent to Sowerby by Dr. Gilchrist.
Probably it is correct to say that less than a dozen complete specimens are
known (unless some have since been obtained by private collectors).
Tomlin and Shackleford, to whom two juveniles were submitted some
years ago, remarked on their ‘extreme likeness to some of the forms from the
Barton Beds and the Paris basin’; in fact they doubted whether the shells were
really South African!
Smith stated that the Mauritian delessertiana has at least 15 columella
pleats, but, as in the case of ponsonbyz, made the risky statement that queketti was
a smaller species, and again without giving the dimensions of the protoconchs.
The largest known specimen is 14. mm. longer than the type, and may be more
comparable with delessertiana in size.
The 51 mm. specimen fortunately retains the operculum, showing that
this species is correctly assigned to the subgenus Lyria. When the animal was
removed is not known.
Named after Mr. Quekett, then curator of the Durban Museum.
Voluta bullata Swainson
‘Fig. 6
1829. Swainson. ool. Illustr., ser. 2, vol. i, pl. 15 (Voluta, pl. 1.).
1859. Chenu. Man. Conchyl., i, fig. 956.
1901. Smith. loc. cit., p. 234.
Spire low, blunt; aperture 4 times spire (profile from suture to apex; 5-6
times if true vertical height measured), occasionally 44 or even 5 times. Proto-
conch 1 (14) whorls, diam. 4 mm. Postnatal whorls 3 (34); smooth, without
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 23
ribs, shoulder, or spiral striae, except some spiral grooves on lower half of body
whorl. Columella pleats 2, with an obscure third one above; a narrow parietal
callus in the posterior angle of the aperture. 62 X 30 mm.
Operculum and radula unknown.
Pale brown with darker speckling and irregular marks which form three
faint spiral bands, towards the outer lip spiral lines in pairs or threes: juveniles
(16 mm.) with dark spiral line dotted with white. The 1st whorl has a ‘neck-
lace’ of alternating brown and white spots at the suture. Callus dark chestnut-
brown.
Port Elizabeth (Sowerby); Algoa Bay (Reeve); Port Alfred (Bartsch,
Turton).
Fic. 6.
Voluta bullata Swainson. Slightly oblique apical views of protoconch of 16 mm. shell (left) and
29 mm. shell (right).
Port Elizabeth and Algoa Bay, St. Francis Bay (= Jeffreys Bay), Still Bay;
all dead and more or less worn (S. Afr. Mus.).
Not taken by the Valdivia or the Pieter Faure. No record of a living specimen,
though dead shells are fairly common.
Remarks. The smallest specimen in S$. Afr. Mus. is 16 mm. long. Measure-
ments of other (selected) specimens are: 55 X 27,59 X 29,57 X 25, 60 X 29,
Ga 20, 62 X 30.
The colour of the parietal callus seems to fade more rapidly than the
external markings of the shell, although not so exposed to abrasion.
Type in the British Museum.
Subgen. Afrivoluta Tomlin
1947. Tomlin. 7. Conch., xxii, p. 244.
Four strong columella pleats. Operculum and animal unknown.*
Proposed by Tomlin as a genus, but here treated as subgenus.
* In an American dealer’s catalogue (1957) this species was illustrated, and a specimen
“Taken alive and perfect’ was offered for sale. Presumably the animal was extracted and thrown
away.
24. ANNALS OF THE SOUTH AFRICAN MUSEUM
Voluta (Afrivoluta) pringler Tomlin
1925. Thiele. D. Tiefsee Exp., xvii, p. 199, pl. 34 (22), fig. 18 (Voluta sp.).
1947. Tomlin. loc. cit., p. 245, text-fig.
Spire low, blunt; aperture 34 (juv.) to nearly 3 (adult) times spire.
Protoconch 2 whorls, diam. 7:5-8 mm. Postnatal whorls 3; smooth, with fine
lines of growth, very fine spiral striae in the type, but none visible in the S. Afr.
Mus. adult or juveniles. A flat sutural band of callus adnate to the previous
whorl, with irregular upper margin, on body whorl expanding into the large
oval or subcircular parietal callus (c. 30 mm. diam.) extending from upper
suture on body whorl to below the posterior end of aperture; this callus is the
easiest means of distinguishing the start of the Ist postnatal whorl from the
protoconch. Columella pleats 4, very prominent; columella projecting slightly
below base of aperture. Outer lip in adult slightly exsert, and reflexed in lower
half 44 x 14in. (¢. 110 X 40 mm.) (Tomlin); 43 x 144 m. (20 x 4gpmann.
(S. Afr. Mus.); juveniles 34-35 X 15-16 mm. (S. Afr. Mus.).
Chestnut-brown with 2 broad bands of pale reddish brown on body
whorl, sutural edging white, columella pleats red, interior rusty reddish-brown,
callus reddish (Tomlin). The S. Afr. Mus. adult is similar, but the general
colour, including columellar pleats and aperture is orange-salmon, the bands
faintly whitish, callus white.
35° 11'S. 23° 2’ E., 500 metres (Thiele); SE. of Cape Recife, 120 fathoms,
and off Jeffreys Bay (St. Francis Bay), west of Cape Recife (Tomlin).
Off Glendower Beacon (Port Alfred area), 100 fathoms; 34° 27'S.
25° 42’ E., 250 fathoms; and off Cape St. Francis, 75 fathoms; one juvenile
from each station (S. Afr. Mus. P.F. coll.).
The Valdivia locality is farther west on the Agulhas Bank than the other
localities. The single broken specimen came up in the same haul with a living
Neptuneopsis gilchrisiz; it measured 62 < 24 mm., but owing to its fragmentary
condition Thiele refrained from naming it.
The Pieter Faure obtained only the three juveniles recorded above.
Remarks. As ‘Tomlin said this is the most noteworthy South African shell
discovered since the Cape Government trawler Pieter Faure obtained Neptune-
opsis gilchristt in 1897; but he forgot Pleurotomaria africana which was discovered
by the Fisheries Survey vessel Africana in 1931 but noc described and named
until 1948. |
Gen. VoLuTocorsis Dall
1890. Dall. Trans. Wagner Free Inst., iii, p. 74. (type t limopsis Conrad).
1929. ‘Thiele. Handbuch Syst. Weicht., i, p. 344 (s.s. type abyssicola).
Operculum absent. Columella with pleats. Radula with tricuspid central
tooth, and a transversely oblong, unicuspid lateral tooth.
w
i.
‘ie
1
4
4
®
»
b
x
if
:
*
§
a)
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 25
Remarks. See Sowerby (loc. cit. infra). Dall included fossil and Recent
species, but Thiele restricted the genus to contain two Recent species only.
Allied to the fossil Volutilithes Swainson 1840 (type f spinosa Lam.).
Volutocorbis abyssicola (Ad. & Rve.)
Figs. 7(a), 9(a)
1848. Adams & Reeve. ool. Samarang. Moll., p. 25, pl. 7, fig. 6, juv. (Voluta a.).
1886. Watson. Rep. Challenger, xv, p. 258, pl. 15, fig. 1, adult (Voluta a.).
1889. Studer. Forschungsreise Gazelle, iii, pp. 52, 55-
1900. Woodward. Proc. Mal. Soc., iv, p. 121, pl. 10, figs. 4-8, 10, 12 (anatomy).
1901. Smith. loc. cit., p. 235 (Volutilithes a.).
1902. Sowerby. Mar. Tacs. Seems Ls P- 97; pl. 2, fig. 6 (radula) enareny) (Volutilithes a.).
1903. Von Martens. D. Tiefsee Exp., vii, p. 31 Gieanlee G3);
nage iihrele. ibid., p. 170; pl. 9 (4), ae 65 (radula).
1922. Cooke. loc. cit., p. 7 (radula).
1925. Thiele. D. Tiefsee Exp., xvii, p. 200.
Numerous fine axial ribs and spiral lirae forming a cancellate sculpture,
with sharp points at the intersections, the first spiral lira with its points forming
a narrow shoulder but not concealing the suture; intervals between Ist and
end lirae (sometimes between 2nd and 3rd) greater than intervals between the
following lirae, forming a shallow groove a little distance below the suture;
axial ribs 18-21 on Ist whorl, increasing to 60 or 70 (80 in largest specimen)
on body whorl. Aperture 2% (juv. 14 mm.) decreasing to 24 or 2 times spire.
Protoconch 2 (24) whorls, diam. 2 mm. Postnatal whorls 5. Columella pleats
I in protoconch, 2 in 1st whorl, increasing to 10-12 in adult, but very variable
in adults, some pleats being smaller intermediaries between pairs of larger
pleats, being as Thiele stated (1925, p. 200) the spiral lirae of the previous
whorl overlaid by callus; he regards only the 4 lowermost as true columella
\
SS
QD
SS
a
6 c
Fic. 7.
(a) Volutocorbis abyssicola (Ad. & Rve.) variation in columella pleats in adults. (b) Fusivoluta
pyrrhostoma (Watson) protoconch of typical form. (c) worn protoconch of forma major.
26 ANNALS OF THE SOUTH AFRICAN MUSEUM
pleats. From the columella a thin parietal glaze extends over base, not con-
cealing the underlying sculpture, its edge adnate to the shell but sometimes
with a very slight free edge at the end of the rostrum. Outer lip in adult slightly
exsert, margin denticulate, internally plicate. 3$ x 14 in. (96 X 38 mm.)
(Watson). A specimen in S. Afr. Mus. has same width and was probably of
equal length (protoconch and end of canal broken). 97 X 40 mm. (in a
private collection).
Smallest specimen in S. Afr. Mus. 11-5 X 6 mm., protoconeh plus 2
whorls. Smallest specimens with denticulate outer lip, presumably mature,
AS x 23 and 46 < 21 mm.
Pale horny or biscuit-coloured, glossy when fresh; some half-grown
specimens show 4-5 faint bands of slightly darker spots; interior of aperture
faintly pinkish or orange. As preserved, animal dark; in life it might possibly
be mauve or purplish (cf. lutosa, infra).
Radula with 100-105 rows; the oblique hind margins of the lateral teeth —
are sometimes slightly crenulate.
Off Cape of Good Hope (Adams & Reeve, Watson); 34° 6’ S. 18° 6’ E.,
117 fathoms (Studer); 34° 43’ S. 18° 30’ E., 125 fathoms (Sowerby) ; 33° 41 S.
18° o’ E., 178 metres; 35° 16'S. 22° 26’ E., 155 metres; and 94° 93) \snume am
318 metres (von Martens).
30° 2'8. 15° 2’ E., 409 fathoms (s.s. Africana, per UGyiae
Numerous stations along the south-western slope of the continental shelf
off Cape Point and the west coast of the Cape Peninsula, 85-230 fathoms, live
and dead specimens; also dead shells from the Agulhas Bank and its southern
margin (vide infra) (S. Afr. Mus. P.F. coll.).
Remarks. Plump and slender individuals occur: e.g. 70 X 33 and 68 x 28
mm. The protoconch is almost always more or less corroded.
Typically the axial ribs and spiral lirae are approximately equally strong.
But there is a tendency for the spiral lirae to become weaker or even obsolete,
thus producing a ribbed instead of a cancellate sculpture. This is well shown in
a series of I1 specimens (24-38 mm. long) containing both cancellate and
ribbed examples from Brown’s Bank, approx. 364° S. 21° E., 100-200 fathoms.
The first row of prickles near the suture, and the 2nd row are distinct (and
sometimes the 3rd), but across the middle of the whorl they disappear, leaving
the axial ribs smooth; the spiral lirae reappear towards the base (P.F. coll.).
Three ribbed examples, 27-36 mm. long., were also taken off Cape Seal,
8o fathoms, and off Cape St. Blaize, 85-90 fathoms (P.F. coll.).
Two dead examples (one of them given to Tomlin) from 34° 34'S.
18° 32’ E., 100 fathoms, 60 x 29 mm.: sharp cancellate sculpture but spiral
lirae on body whorl tending to greater prominence than the axial ribs: proto-
conch narrower than in typical examples, diam. 1:5 mm. (slightly corroded) ;
columella pleats 2 with one intermediary and 3 above very feeble; parietal
glaze adnate to shell without free edge; outer lip broken away. In some
respects approaching ludosa (P.F. coll.).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 27
Four living and 2 dead examples, 31 xX 16 to 35 X 17 mm. long, from
off South Head, Saldanha Bay, 190 fathoms: protoconch diam. 2 mm.; axial
ribs predominating, sculpture in the youngest sharp, in the others more or less
corroded; columella pleats 2 plus 4 small (in the youngest), 1 plus 6 or 7, the
latter decreasing in size posteriorly (2 specimens), 2 plus 1 intermediary and
3 small above (2 specimens); parietal glaze with free edge except in the
youngest; outer lip except in the youngest thickened by close aggregation of
growth-lines externally, internally feebly denticulate and plicate in one speci-
men, with obscure traces of denticles in two others. Radula as in abyssicola
(PF. coll.).
In having a free edge to the parietal glaze these specimens also approach
lutosa.
One living and one dead from off Duminy Point, 87 fathoms (slightly
farther north than South Head), 45 x 22 (living), 50 x 24 mm. (dead): apex
‘acute’, protoconch narrow, diam. 1°5 mm.; cancellate sculpture sharp on
upper whorls, somewhat corroded on body whorl; columella pleats 3, upper
one small, plus 2 obscure (smaller specimen), 2 plus 1 intermediary and 1 above
obscure (larger specimen); parietal glaze with free edge; outer lip broken.
Radula as in abyssicola (P.F. coll.).
These shells appear to be referable to lutosa.
var. lutosa Koch
ro4G. Koch. jf. Conch., xxiti, p. 5, pl. 2.
Similar to abyssicola. Koch said ‘apex acute’, but the acuteness appears
due to corrosion; sculpturing not so sharp, which is also due to corrosion,
young specimens being like abyssicola; columella pleats 3-5, the upper 2-3 being
feeble; in larger examples the parietal glaze thicker, nearly concealing the
sculpture on body wall, because the animal deposits the glaze over the layer of
clay covering the shell, consequently when the shell is cleaned the edge of the
glaze is free not adnate. 80 X 35 mm. (Koch).
Cream (when not corroded), interior of aperture pale orange-brown,
animal mauve (Koch).
Off Port Nolloth and Orange River mouth, 40-60 fathoms (Koch).*
32° 9' 8. 18° 6’ E., 59 fathoms (s.s. Africana, per U.C.T.). Frequently encased
in stiff red-brown or umber-brown clay.
Remarks. The P.F. examples described above as variants of typical abyssicola
impair the validity of this form as a full species. I regard it, at most, as a
variety. The two forms seem to overlap in the vicinity of Saldanha Bay, and
the slight differences appear to be due to habitat.
I have seen 15 Africana examples (AFR. 718 B.) from 26 to 75 mm. long.
* In the same American dealer’s catalogue mentioned in the footnote, p. 23, a ‘Paratype’
was offered for sale.
28 ANNALS OF THE SOUTH AFRICAN MUSEUM
Volutocorbis gilchristi (Sow.)
1902. Sowerby. Mar. Invest. S. Afr., ii, p. 99, pl. 2, fig. 5.
Spire high, aperture 13 times spire. Protoconch diam. 1-5 mm. (broken).
Postnatal whorls 4; axial ribs c. 22 on 1st whorl, ¢. 24 on body whorl
(Sowerby: 16, ? typ. err. for 26); spiral lirae obscure on 1st—3rd whorls, but
distinct on body whorl, especially towards base; top of whorl projecting above
the sunken suture as a crenulate or denticulate ridge, somewhat corroded and
not so noticeable on upper whorls; 2nd spiral lira separated from ridge farther
than the following lirae one from another. Columella pleats 6, decreasing in
size posteriorly. Outer lip reflexed, thickened, several closely aggregated
growth-lines forming a stout varix externally, internally obscurely denticulate.
Parietal glaze slight, adnate to shell. 30 x 15 mm. (Sowerby); 28 X 13 mm.
(cotype, S. Afr. Mus.).
White with pale cream glassy periostracum.
Off Cape Natal (Durban), 200 fathoms (Sowerby).
Same locality, 185-200 fathoms (S. Afr. Mus. P.F. coll.).
Type ? in coll. Sowerby (? Brit. Mus.); cotype in 8S. Afr. Mus.
Remarks. Distinguished by the deeply sunken suture. Von Martens when
describing (1903) epzgona from East Africa did not mention gilchristi; and
Thiele in his Handbuch (1929, p. 345) seems to recognize abyssicola and epigona
as the only two species in the genus.
V. epigona (1903. D. Tiefsee Exp., vii, p. 106, text-fig.) is slightly larger than
gilchristi (text said 30 mm., but the line alongside the enlarged figure measures
33 mm.). It differs in having 8 columella pleats, the upper ones much more
strongly developed than in gilchristi; and strong plicae within the outer lip
which is not thickened. The suture is not mentioned, but judging by the figure,
is not so deeply sunken. 3
Gen. FULGORARIA Schumacher
Columella with 6-8 pleats.
Saotomea Habe 1943 was described as a section or subgenus with one
columella pleat and a lozenge-shaped operculum.
Fulgoraria blaizei n. sp.
Fig. 8(b)
Aperture subequal to spire. Protoconch mammiliform, 2 (24) whorls,
diam. 2°8-3:5 mm., smooth, junction with rst postnatal whorl indistinct in
three specimens but abrupt in the 24 mm. specimen. Postnatal whorls 4,
profile evenly curved, without shoulder; slightly arcuate low axial ribs 16-18
on Ist whorl, 18-20 on and, ill-defined and petering out on 3rd whorl; fine
spiral lirae traceable only on ist and 2nd whorls. Growth-lines fine, arcuate.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 29
About 20 spiral lirae on base. Aperture narrow ovate, canal rather short.
Columella with one pleat, best seen on the 39 mm. specimen (taken alive),
very indistinct on the dead ones. 42 X 15 mm., 39 X I1I°5mm., 24 X 9°5mm.,
18 X 6-5 mm.
Operculum obovate, nucleus apical (but apex broken), 8-5 xX 3:5 mm.
in 39 mm. shell with aperture 17 mm.
Uniform salmon-buff, protoconch white, operculum pale amber; the two
dead specimens pale buff.
Off Cape St. Blaize, 73 miles, 125 fathoms, one 42 mm., one 24 mm.,
and one 18 mm., dead; same locality, 105 fathoms, one living 39 mm. (S. Afr.
Mus. A3433 type (live), A3430 cotypes. P.F. coll.).
Remarks. Provisionally placed in the genus Fulgoraria.
There is no record whether these specimens were submitted to Sowerby;
probably not. Nor is it known when the animal was removed from the only
specimen taken alive.
} The spiral lirae are finer and closer together in the 42 mm. and 24 mm.
specimens, than in the type and the smallest specimen.
Gen. FustvoLuTa von Martens
1902. Von Martens. SB. Ges. naturf. Fr. Berlin, p. 237.
Operculum oval, nucleus apical, curved to left. Columella without pleats.
Radula with tricuspid central tooth, no lateral.
Fuswoluta pyrrhostoma (Watson)
Figs. 7(0), 9(4)
1882. Watson. 7. Linn. Soc. Lond., xvi, p. 374 (Fusus (Sipho) p.).
1886. id. Challenger Rep., xv, p. 208, pl. 12, fig. 2 (Fusus (Sipho) p.).
? 1889. Studer. Forschungsreise Gazelle, ii, pp. 52, 54 (Fusus mandarinus, non Duclos).
1903. Sowerby. Mar. Invest. S. Afr., lu, p. 226, pl. 3, fig. 1 (shell, operculum, radula,
anatomy).
1903. Von Martens. D. Tiefsee Exp., vii, p. 32, pl. 3, fig. 15.
? 1903. id. ibid., p. 33 (Fusus mandarinus, non Duclos).
1903. Thiele. ibid., p. 171, pl. 9 (4), fig. 67 (radula).
Aperture subequal to spire. Protoconch 23 whorls, diam. 2-5 mm. Post-
natal whorls 5; axial ribs on ist whorl 13-14, on body whorl 15-20, but
gradually becoming obsolete, arcuate, beginning at suture and sometimes
forming a very slight shoulder, obsolete on base, sharp (when not worn),
narrower than intervals; crossed by ¢c. 15-18 or 20 spiral lirae. Columella
slightly curved, parietal glaze concealing sculpture, adnate to shell without
free edge. 42 X 17 mm. (Challenger: 38 mm.).
Operculum 12 X 5 mm. in 42 mm. shell.
Dull white, fresh specimens with very thin pale fawn periostracum,
aperture internally pale orange-salmon.
30 ANNALS OF THE SOUTH AFRICAN MUSEUM
Radula 1:5 mm. long, with c. 45 rows. Thiele’s specimen was also 1-5 mm.
long.
34° 41'S. 18°36’ E., 98 fathoms (Watson); 33° 59'S. 17° 52’ E., 50
fathoms (Studer); 34° 20’ S. 18° 36’ E., 70 metres, and 34° 33°52) aa iE.
318 metres (von Martens). Sowerby gave no locality.
Mouth of False Bay and off Cape Point and the west coast of the Cape
Peninsula, 45-200 fathoms; Brown’s Bank (approx. 364° S. 21° E.), 80-100
fathoms; Cape St. Blaize, distant 73 miles, 125 fathoms (S. Afr. Mus. P.F.
coll.).
Remarks. It is most unlikely that the New Zealand Fusus mandarinus occurs
off Cape Point. Moreover von Martens’s characterization of the two dead.
Gazelle shells might well apply to some of the present examples: superficially
(‘aiisserlich’) similar [to pyrrhostoma] with narrower (‘feinere’) apex unlike that
of a Voluta, ribs disappearing on 6th [i.e. penultimate or 4th postnatal whorl]
and relatively greater length of the visible portion of the whorls; 31 mm. long.
Except for the last, rather vague, character there are similar worn shells in the
P.F. series with narrow pointed protoconch.*
The columella is usually more curved than in Watson’s figure.
forma major n.
Figs. 7(¢), 9(¢)
Two specimens considered by E. A. Smith and L. J. Shackleford to be
extra large specimens of pyrrhostoma.
Off South Head (Saldanha Bay), 190 fathoms. 60 X 24 mm., smaller
specimen 55 X 22 mm. with protoconch and operculum, taken alive but
animal not preserved (A3420).
Both specimens much corroded, even the live one. Traces of axial ribs
remain on the 3rd and 4th whorls, but apparently they were not developed
on the body whorl, thus resembling the typical form (S. Afr. Mus. P.F. coll.).
A live specimen, 74 X 25 mm., was taken by the s.s. Africana (AFR. 738
D., submitted per U.C.T.) at 30° 21’ S. 16°50’ E., 185 metres. Protoconch
and upper whorls corroded. Postnatal whorls 6; axial ribs well marked but
feeble and irregular on back of last whorl and outer lip; spiral lirae obsolete.
Radula 5 mm. long, with 58 teeth.
Fuswwoluta capensis (Thiele)
1925. Thiele. D. Tiefsee Exp., xvii, p. 179, pl. 31 (19), fig. 27 (Glypeuthria ? c.).
1931. Tomlin. Ann. S. Afr. Mus., xxx, p. 165, fig. 6 (Glypeuthria capensis, non Thiele).
1945. id. J. Conch., xxii, p. 135 (Glypeuthria sculpturata nom. nov. for capensis Tomlin preocc.).
1957. Barnard. ibid., xxiv, p. 210 (radula, generic position).
* Krauss (1848, p. 110) mentions the similarity of his Fasciolaria badia (= lugubris Rve.) to
Fusus mandarinus; the Gazelle shells may be this species, but I think they are far more likely to
be pyrrhostoma.
So
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 31
Aperture slightly shorter than spire (by about the length of the proto-
conch). Protoconch 2 whorls, diam. 2 mm. (but corroded). Postnatal whorls
6. Axial ribs 12-14 on 1st whorl (but this whorl usually corroded), 18 on body
whorl, slightly arcuate, beginning at suture, extending across base; crossed by
18-20 strong spiral lirae which are finer in the upper part of the whorl near
suture. Columella curved, canal short; parietal glaze concealing sculpture,
adnate, without (sometimes very slight) free edge. 38 K 15 mm.
Operculum oval, nucleus apical (but usually broken), curved to left,
7 X 4mm. in 29 mm. shell.
Greyish white, operculum amber coloured.
Radula 2-75 mm. long, with 45-50 teeth, indistinguishable from that of
pyrrhostoma.
35° 9’ S. 18° 33’ E., 564 metres (Thiele) ; off Cape Point, 318-400 fathoms
(Tomlin).
In addition to those sent to Tomlin, there are specimens from between 250
and 560 fathoms off Cape Point (S. Afr. Mus. P.F. coll.).
Remarks. Comparison of the smallest P.F. specimen, although twice as large
as Thiele’s Valdivia specimen, with his figure leaves no doubt as to the identity.
Tomlin did not see this specrmen. Nor was any animal sent to Tomlin.
This species appears to live in deeper water than pyrrhostoma.
Though the two species have a slightly different appearance—capensis is
less fusiform and more distinctly spirally lirate—the descriptions read very
much alike as regards details. Possibly intergrading forms will be found.
Fusivoluta decussata n. sp.
Fig. 8(c)
Aperture (as preserved) subequal to spire without protoconch. Proto-
conch mammiliform, 2 (24) whorls, diam. 3-5 mm., smooth but with a few
axial pliculae towards the junction with Ist postnatal whorl where the spiral
lirae start. Postnatal whorls (as preserved) 4, profile evenly convex; with
slightly arcuate axial ribs and spiral lirae producing a cancellate sculpture, the
ribs a little more prominent than the lirae, intersections slightly nodulose;
¢. 30 ribs on Ist whorl, ¢. 45 on and, 55 on last whorl (but some feebler and
closer together than others making an exact count difficult); 7 lirae on Ist
whorl, 8 on 2nd, g on 3rd, and to on last whorl, about 6 additional on base.
Columella slightly curved, without pleat. Aperture oval, canal narrow (but
lip broken). 35 X 12 mm.
Buffalo River (East London), 15 miles, 310 fathoms, one dead specimen
and one fragment of a juv. (S. Afr. Mus. A3432. P.F. coll.).
Remarks. This distinctive shell is placed provisionally in Fusivoluta, although
the animal is unknown. The protoconch is larger than in fyrrhostoma, and
relatively larger than in anomala von Martens,
32 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fusivoluta elegans n. sp.
Fig 8(a)
Aperture a little longer than spire (11 : 8 mm.). Protoconch 14-2 whorls,
alt. 1 mm., diam. 1:2 mm., smooth, with a few axial pliculae towards junction
with 1st postnatal whorl. Postnatal whorls 44-5, profile convex, on the lower
whorls the upper part flat but without definite shoulder. Axial ribs 13 or 14
on 1st whorl, 15 on 2nd, 16 on grd, 17 on 4th, and 19 on last whorl; on
penultimate and last whorl the growth-lines become prominent and form
intermediary riblets, usually 2 (3) between each pair of main ribs; ribs obsolete
on base; crossed by spiral lirae 5 on 1st whorl, 7 on 2nd, 9 on 3rd, 12 on 4th
and 13-14 on last whorl; on 3rd—5th whorls lirae slightly stronger on the lower
convex part than on the upper flat part of whorl, intersections with main ribs
slightly nodulose, those with the intermediaries a little more conspicuous, ¢. 20
additional lirae on base. Columella almost straight, without any indication of
a pleat. Canal rather long. 19 X 6:5 mm.
Off East London, 400 fathoms, one dead (S. Afr. Mus. A8803. P.F. coll.).
Remarks. Placed in Fusivoluta because the sculpturing is essentially similar
to that of pyrrhostoma, anomala and capensis. ‘The size of the protoconch (unworn),
however, indicates a smaller species.
Somewhat resembling in shape /ilgendorfi von Martens (1897) placed in
the genus Benthovoluta Kuroda & Habe (1950. Illustr. Cat. Jap. Sh., no. 5).
Fic. 8.
(a) Fuswoluta elegans n. sp. (b) Fulgoraria blaizei n. sp., with protoconch further enlarged.
(¢) Fusivoluta decussata n. sp., with protoconch further enlarged,
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 33
Gen. NEPTUNEOPSIS Sow.
1898. Sowerby. Mar. Invest. S. Afr., i, p. 5 (exact date ? printer’s order number on cover
of reprints gives . . . 3, 1898, which presumably means March).
1900. Woodward. Proc. Mal. Soc., iv, p. 120 (anatomy).
1902. Pace. ibid., v, p. 25 (anatomy).
Columella without pleats. Operculum ovoid, nucleus apical. Radula with
tricuspid central tooth and degenerate lateral.
The genus is monotypic.
Neptuneopsis gilchristt Sow.
Fig. 9(d)
1898. Sowerby. loc. cit., p. 6, pl. 1 (shell, operculum, radula).
1900. Woodward. loc. cit., p. 120, pl. 10, figs. 2, 3, 11, 13, 14.
1902. Pace. loc. cit., p. 25, pl. 2, figs. 5-8.
1903. Von Martens. D. Tiefsee Exp., vii, p. 33.
1903. Thiele. ibid., p. 171, pl. 9 (4), fig. 68 (radula).
Protoconch conical, apex pointed, usually slightly lop-sided, size variable:
meee 7, 10 X 6-5, 11 X 7, 11 X 8-5 mm. Postnatal whorls 6; finely
striated axially and spirally, growth-lines coarser, surface dull. Aperture sub-
equal to spire. Outer lip slightly reflexed. Up to 198 x 70 mm.
Shell pinky-white, periostracum amber-brown or greyish-brown, or
slightly olivaceous, operculum chestnut-brown.
Radula: see p. 35.
Off ‘Cape of Good Hope’, 33 fathoms [34° 17’ 8. 18° 35’ E., False Bay]
living (Sowerby); 35° 10’ S. 23° 2’ E., 500 metres, living (von Martens).
Off west coast of Cape Peninsula, 160 fathoms, living (U.C.T. ex trawler).
Off west coast of Cape Peninsula, 60-91 fathoms, living (S. Afr. Mus.
Pe coll.).
Agulhas Bank: off Cape St. Blaize and Flesh Point, 33-105 fathoms, dead;
off Nanquas Peak (eastern part of Algoa Bay), 49-59 fathoms, dead (S. Afr.
Mus- P.F. coll.).
S. Afr. Mus. also has two specimens with opercula from the Ross-Frames
collection, without record of how or where obtained, but probably purchased
from trawlers.
Remarks. The most notable of the many novelties obtained by the Pieter
Faure.
In the Report of the Government Biologist for 1897, p. 10, there is no
mention of the capture of this large mollusc, nor in fact of any of the captures
except fishes.
There was formerly (about 1917) in the Museum of the Zoology Depart-
ment of the South African College (where Dr. Gilchrist was Professor of
Zoology) a dry shell labelled ‘Type’. This may possibly have been the Type,
34 ANNALS OF THE SOUTH AFRICAN MUSEUM
but it appears to have been lost when the Zoology Department moved out to
the new University buildings in the suburbs.
A specimen, shell and animal in alcohol in S. Afr. Mus., was taken on
8 September 1897 at Station IX in False Bay (34° 17’ S. 18° 35’ E.) in 33-40
fathoms, bottom fine white sand. The shell is 170 mm. in length; the operculum
has not been removed from the animal, which has not been dissected. This
specimen therefore cannot be the Type.
Only two other living specimens were taken, both off the west coast of the
Cape Peninsula, one in March 1900, the other in August 1903 (both in S. Afr.
Mus., from one of which I have removed the radula).*
d c
Fic. 9.
(a) Volutocorbis abyssicola (Ad. & Rve.), central and lateral plates of radula. (b) Fusivoluta pyrrho-
stoma (Watson), central plate of radula. (c) F. pyrrhostoma forma major, central plate of radula.
(d) Neptuneopsis gilchristi Sow., three central plates with degenerate lateral plates from near front
end of radula; the expansion of the basal membrane thins out laterally as indicated by the
spacing of the dots.
It seems therefore that ‘wo living specimens were taken at Station IX in
False Bay. One sent to Sowerby and described by him in 1898; he himself
seems to have extracted the radula and passed the animal (thus mutilated) on
* Pace (loc. cit., p. 21 and p. 25 footnote 2) records that the British Museum secured a
spirit specimen (reg. no. 1901: 10.29.10) in 1901, which he dissected. There seems to be no
record in the Pieter Haure log-book of the capture of this specimen.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA »} 35
to Woodward for dissection (see Woodward 1900, and Pace 1902). The second
specimen is that now in S. Afr. Mus.; it can be regarded as no more than a
topotype.
The Valdivia obtained a living example at 500 metres on the southern
slope of the Agulhas Bank, indicating that the habitat probably extends west-
wards along the slope of the continental shelf to Cape Point and along the
western coast of the Cape Peninsula. The locality at the mouth of False Bay
may be regarded as an outlier, or possibly an exceptional occurrence.
The Pieter Faure, whose primary object was the discovery of inshore com-
mercial fishing grounds, scarcely touched the southern fringe of the Agulhas
Bank; which probably explains why only dead shells were obtained, except on
the three occasions mentioned.
These dead shells came from two areas: one south of Flesh Point and Cape
St. Blaize, the other at the eastern end of Algoa Bay. These two ‘pockets’ seem
to indicate bottom currents flowing inshore, either westwards or possibly as
reverse compensatory currents eastwards. As yet it is impossible to say whether
the shells from the Algoa Bay pocket indicate a habitat of the living mollusc
farther east on the continental shelf towards East London.
Radula (fig. 9d). Pace (loc. cit., p. 27 footnote 1) said, ‘Sowerby’s figure
shows such discrepancies from my preparation that I venture to question
whether the radula of . . . e.g. Cymbiola ancilla, may not have been accidentally
substituted .. .’. This statement is quite unjustified. Pace’s own figure (pl. 2,
fig. 8) is quite comparable with those of Sowerby and Thiele (except for the
inclusion of the lateral teeth), but none of these three figures is comparable
with Pace’s figure of C’. ancilla (pl. 2, fig. 9); in the former the notches between
the cusps are U-shaped, in the latter V-shaped.
In the specimen examined by me the teeth are closer together than in
Sowerby’s and Thiele’s figures, more as drawn by Pace.
Pace (p. 28) said the lateral teeth disintegrated so rapidly in KOH that
he had no time to make a drawing. He is perfectly correct in recording their
presence. They can, however, scarcely be called teeth as they have no free
margins; they are in fact indicated only by oblique thickenings in the basal
membrane. They correspond in number with the central teeth, and are
undoubtedly degenerate lateral teeth. There are 70-75 rows.
Fam. HARPIDAE
1916. Melvill. 7. Conch., xv, pp. 25-40.
1939. Peile. Proc. Mal. Soc., xxiii, p. 271, fig. (radula).
The only specimen of this family obtained by the P.F. was a 45 mm.
H. conoidalis Lam. (identified by Sowerby), from off Umvoti River (Natal),
27 fathoms.
Also obtained at Delagoa Bay (U.W.).
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fam. VASIDAE
Bartsch (1915, p. 42) records Xancus globulus Chemn. from “Cape of Good
Hope’. This is certainly not an indigenous South African species.
Vasum turbinellum (Linn.) is also doubtfully indigenous, though it occurs
at Mozambique Island (U.W.).
Turton’s ‘Xancus sp.’ (1932. p. 45, pl. 10, no. 333), 3°5 X 2 mm., appears
to be a juvenile Mitra.
Vasum truncatum Sow.
Fig. 10
1892. Sowerby. Mar. Sh. S. Afr., p. 17, pl. 4, fig. 85 (adult, worn).
1902. Smith. 7. Conch., x, p. 249, pl. 4, fig. 6 (triangularis, juv.).
1903. id. Proc. Mal. Soc., v, p. 370, pl. 15, fig. 3 (adult).
Shell thick in adult, conical, spire very low but in juvenile protoconch and
first whorl (or first two whorls) forming a projecting papilla. Protoconch: »v.
infra. Postnatal whorls 6. Shoulder with c. 11 blunt knobs, whose inclusion in
succeeding whorls produces an undulate
suture. Base with 4 or 5 blunt spiral
ridges, more or less interrupted and
nodose. Columella with 4 pleats, more or
less equally strong; rimate anteriorly at
snout but umbilicus open. Periostracum
thick, scabrous with close-set growth-lines.
65 X 50 mm. (Sowerby); 48 X 35
mm. (Smith); 66 x 55 mm. (S. Afr. Mus.
beach-worn, protoconch and first 3 or 4
whorls missing and end of canal worn);
41 (protoconch missing) X 32 mm., and
55 X 43 mm. (width across shoulder
knobs) (S. Afr. Mus., .the latter specimen
retains most of the periostracum).
Port Elizabeth, Port Alfred (Sowerby,
Bartsch, Turton); Port St. Johns (S. Afr.
Mus.). In deep water off Durban and
Port Shepstone (Smith); off Durban (S.
Fic. 10. Afr. Mus.).
Vasum truncatum Sow., protoconch and WN juvenile obtained by the P.F. off
first two postnatal whorls of juvenile : ;
15 X 10mm. Umkomaas River (Natal), 40 fathoms, is
worth a separate description and figure.
15 X 10 mm. Protoconch 24~3 whorls, papilliform, gradually passing
into first postnatal whorl without distinct junction; 1st whorl somewhat worn,
and with 5 or 6 very faint spiral carinulae, superseded at 24 whorl by faint and
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA | 37
irregularly spaced axial pliculae, c. 16 in three-quarters of a whorl (may be
either protoconch or Ist postnatal whorl). Postnatal whorls 2, the axial pliculae
superseded by 6 spiral lirae, the lower 2 more prominent and forming a double
peripheral keel. On 2nd whorl c. 8 lirae above the keel, below the keel c. 12
additional lirae with a weaker intermediary between each pair. Faint spaced
growth-lines are traceable, but no axial ribs; the circumference, however, when
seen from the apex is gently undulate, c. 11-12 lobes on 1st and 2nd whorls;
these undulations are not prominent enough to cause nodulose projections on
the base as they do in larger examples. Columella pleats 4, the 3rd thinner
than the others (as is the case in two larger specimens 41 and 55 mm. in length).
Snout slightly rimate.
Remarks. Type of truncatum in Bairstow collection (Oxford Mus.); of
triangularis in British Museum.
Smith (1903) gave an emended description and recognized his ériangularis
as the not fully adult of truncatum; but he did not give the size of his Port
Shepstone adult.
Fam. MITRIDAE
1919. Cooke. Proc. Zool. Soc. Lond., p. 405 (classification according to radulae).
1922. Peile. Proc. Malac. Soc., xv, p. 93 (radulae).
1936. id. ibid., xxii, p. 141 (radulae).
1937. id. ibid., xxii, p. 181 (radulae).
Cooke’s paper on the radulae was based on the Gwatkin collection in the
British Museum. Unfortunately, it seems that reliance cannot always be
placed on the correct naming of the slides. Some errors have been detected
(e.g. Euthria queketii). Suspicion also arises in one case in this family, namely
‘circula var.’, under which Cooke (p. 415) said: “There is evidently some con-
fusion in the specimens forwarded [to Gwatkin] by Mr. Burnup....’ But this
did not prevent the description of a new species burnupiana! (see pp. 48-49).
The results contained in Peile’s papers, which were based on radulae
extracted by himself from the respective shells, are far more acceptable.
Gen. Mirra Lam.
More or less fusiform. Central plate of radula more or less quadrate, with
comparatively few (4-8) cusps, lateral plate broader, usually considerably
broader than central, usually with numerous cusps, of which one or two may be
enlarged. Ocasionally both central and lateral plates are unicuspid. The group
containing czrcula has an arcuate central plate with numerous small cusps, the
lateral also with numerous small cusps.
Mitra caffra (Linn.)
1758. Linne. Syst. Nat., ed. 10, p. 732 (Voluta c.).
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 208.
1935. Dautzenberg. Mem. Mus. R. Hist. Nat. Belg., H.S. II, 17, p. 120 (Turricula c.)
(references).
38 ANNALS OF THE SOUTH AFRICAN MUSEUM
Aperture 14-14 times the spire. Postnatal whorls 8 (S. Afr. Mus. speci-
men). Axial ribs 16-18 on early whorls, decreasing to 14-16, and becoming
obsolete on 7th whorl; crossed by 3-4 spiral striae, 5 on body whorl, about 15
additional striae on base becoming stronger anteriorly. Columella pleats 4;
outer lip plicate within. 40 x 15 mm. (S. Afr. Mus. specimen).
Chestnut-brown, with pale yellow or white peripheral band, and a second
band in middle of base and outer lip, base below this band somewhat yellowish.
Delagoa Bay (U.W.); Inhambane (U.C.T.); Mozambique (S. Afr. Mus.).
Distribution. Dar-es-Salaam (S. Afr. Mus.), East Indies, Philippine
Islands, China.
Remarks. ‘Two very worn specimens in the Ross-Frames collection labelled
as from Mozambique.
As ‘Mitra sp.’ Turton (1932. Mar. Sh. Port. Alfred, p. 46, pl. 10, no. 343)
mentions 3 specimens, 26 mm. long, light brown with a darker band at the
sutures. The photograph shows a dark shell with a light band above the suture
and apparently on the upper part of the outer lip. The summit is very rounded,
and. the shell is obviously in an advanced stage of “‘beach-wear’.
_ The resemblance of the photo to caffra is noticeable, and by grinding down
one of the above-mentioned Mozambique specimens a very fair imitation of
Turton’s shell was obtained. There is, however, one difficulty in the way of
this explanation: presumably the summit of Turton’s specimen is solid shell
(the photo is not at all clear), whereas in the artificially produced imitation the
hollow interior of the whorl is exposed.
Turton’s specimens (in spite of their condition!) should be re-examined.
Mitra euzonata Sow.
1900. Sowerby. Proc. Mal. Soc., iv, p. 4, pl. 1, fig. 11.
Axial ribs 11 on 1st whorl, 15-16 on last whorl; spirally punctate-striate
(Sowerby). 10 X 4°5 mm.
White with an orange-brown peripheral band.
Remarks. Very likely merely a colour variety of Vexillum capense. The
suture is no deeper than in the latter.
Mitra kowvieensis Sow.
1901. Sowerby. Proc. Mal. Soc., iv, p. 213, pl. 22, fig. 17.
1932. ‘Turton. Mar. Sh. Port Alfred, p. 47, pl. 10, no. 352 (helena, non Bartsch.).
1932. id. ibid., p. 47, pl. 10, no. 353 (eucosmia).
1933. id. J. Conch., xix, p. 370 (becki nom. nov. pro helena preocc.).
Aperture subequal to spire. Protoconch 14 whorls. Postnatal whorls 4
(Sowerby: 6, i.e. incl. protoconch), profile slightly shouldered. Axial ribs 11
on 1st whorl, 13-14 on last whorl, slightly narrower than intervals, extending
across base; spiral striae 4. on 1st and 2nd whorls, 5-6 on later whorls, crossing
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA - 39
the ribs, on base 4 additional striae (i.e. 5 lirae) followed by 4—5 stronger lirae,
intersections with ribs nodulose. Columella pleats 4. 6 X 2-5 mm.
White; Turton says fresh specimens are slightly pink.
Kowie (Port Alfred).
Remarks. Turton said there were broad and narrow individuals, but in
spite of this he created an. sp. (helena, renamed beckz) ; eucosmia is obviously also
a synonym, although it has fewer spiral striae (cf. capense). Probably synony-
mous with V. capense.
Two dead P.F. specimens appear to correspond with the narrow form
(beckt) of kowieensis: off Tugela River (Natal), 65-80 fathoms and off Umkomaas
(Natal), 40 fathoms (the latter specimen 7-3 x 3 mm.) (S. Afr. Mus. P.F. coll.).
Parmele sspecimen from 30°47 S. 30°29 E., 24 fathoms (U.C.T.),
6 X 2-5 mm., has the protoconch and first two whorls coral pink, the last two
maroon with a single narrow white band (actually a series of spots, one on each
rib) a little below the periphery.
Mitra distincta (Thiele)
1925. Thiele. D. Tiefsee Exp., xvii, p. 186, pl. 32 (20), fig. 25.
Scarcely distinct from kowzeensis. Protoconch 14 whorls, alt. and diam.
0-75 mm. Postnatal whorls 4, profile shouldered. Axial ribs 11 on rst whorl,
16 on last whorl; spiral striae 4 on 3rd, 6—7 on last whorl (incl. 1 or 2 obscure
on shoulder), crossing ribs, 10-11 additional striae on base. 7 X 3 mm.
35° 16’ S. 22° 26’ E., 155 metres (Thiele) ; off Cape St. Blaize, 125 fathoms,
2 dead (S. Afr. Mus. P.F. coll.).
_ Remarks. The ribs become obsolete on the base, not nodulose as in kowve-
ensis. Judging by Thiele’s figure the present specimens have more ribs than the
Valdivia specimens; the latter might be re-examined.
Thiele placed his species in Turricula (now Vexillum) without, however,
having a radula for confirmation.
Mitra bathyraphe Sow.
1900. Sowerby. Proc. Mal. Soc., iv, p. 4, pl. 1, fig. 9.
1932. Turton. Mar. Sh. Port Alfred, p. 46, pl. 10, no. 348 (didyma).
Aperture equal to spire. Protoconch 14 whorls, alt. and diam. 0-5-0-6 mm.
Postnatal whorls 4, profile not shouldered, suture canaliculate, upper edge of
whorls slightly undulate or crenulate. Axial ribs 14 on ist whorl, 16-18 on
last whorl, tending to become obsolete on outer lip, subequal to intervals,
extending across base; spiral striae 6 on 1st whorl, 9-11 on last whorl, about 12
additional striae on base, becoming stronger anteriorly, intersections with ribs
slightly nodulose. Columella pleats 4. 8 X 3:25 mm. Pink.
Kowie (Port Alfred). Off Gove Rock (East London area), 27 fathoms, 3
dead but fresh (S. Afr. Mus. P.F. coll.).
40 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. The P.F. specimens retain the protoconch, and show the spiral
striae crossing the ribs, though they are only feebly impressed on the crest of
the ribs. Like beach examples they are pink in colour.
M. didyma is obviously a very worn example of bathyraphe.
Mitra canaliculata Sow.
1900. Sowerby. Proc. Mal. Soc., iv, p. 4, pl. 1, fig. 10.
Only beach-worn specimens available in 8. Afr. Museum, but they seem
to confirm the absence of spiral striae, which is a distinguishing feature.
Mitra latruncularia Rve.
1844. Reeve. Proc. Zool. Soc. Lond., p. 181; and Conch. Icon., pl. 21, fig. 166.
1932. Turton. Mar. Sh. Port Alfred, p. 45.
Aperture a little shorter than spire (by about length of protoconch and
one whorl). Postnatal whorls 5 (perhaps 6). Spiral grooves crossed by axial
growth-lines which cause the former to appear punctate (especially in worn
specimens), 5 on uppermost whorl, increasing to 7 on body whorl, 13-14
additional grooves on base, becoming deeper anteriorly. Columella pleats 4.
30 X 10 mm.
Purplish- or greyish-brown, with a broad white band at top of whorl and
another from top of aperture around middle of last whorl; the whole with
irregularly scattered orange-brown spots, which may here and there unite into
short axial flames.
Port Elizabeth (Sowerby), Port Alfred (Bartsch, Turton). Jeffreys Bay
and Still Bay (S. Afr. Mus.).*
Remarks. One 30 mm. specimen from Jeffreys Bay has only 10 additional
spiral grooves on the base (i.e. 17 grooves on the outer lip instead of 20-21).
Although only one out of 10 specimens, this shows that variation in the number
of grooves occurs, and form a transition to:
var. albozonata ‘Turton (1932. loc. cit., p. 45, pl. 10, no. 335), which has
3 spiral grooves on the uppermost whorl, increasing to 5 on body whorl, and
7-8 additional grooves on base (i.e. 12-13 on outer lip). Up to 26 mm. long
(apex worn).
Colour as above, but upper white band usually clearer with fewer spots
except actually at the suture, second band not so obvious, sometimes whole
shell brown with a few white spots along the suture.
Port Alfred (Turton); Jeffreys Bay (S. Afr. Mus.).
M. latruncularia and albozonata cannot be separated at a glance by the
coloration, but are easily separated by the number of grooves. M. albozonata
* A set of beach-worn specimens in S. Afr. Mus. is registered as obtained by E. L. Layard
(Curator 1855-72) in “Table Bay’; but this locality record is unacceptable.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 41
can be regarded at most as only a variety. The original description does not
state the number of spiral grooves, and reference to the type is necessary to
show which is f. typica and which the variety.
Mitra picta Rve.
1844. Reeve. Conch. Icon., Mitra, pl. 16, fig. 123.
1903. Von Martens. D. Tiefsee Exp., vii, p. 53.
1932. Turton. Mar. Sh. Port Alfred, p. 46, pl. 10, no. 344 (juv.).
Spire less than aperture (juv. to c. 15 mm.), equal to (c. 20-25 mm.), then
14 (half-grown) to 14 (adult) times the aperture. Protoconch 2 whorls, 2
whorls as hatched alt. 1:75 mm., on later juveniles visible portion alt. and diam.
0:75 mm., smooth. Postnatal whorls 6 or 7 (all adults worn). Spiral punctate
striae 6 on Ist whorl, increasing to 12 on body whorl, 12-13 additional striae
on base (in adult) becoming stronger anteriorly (24-25 on outer lip); fine
close-set growth-lines. Columella pleats 4. 40 X 12 mm.
Castaneous with irregular white marks and flames, sometimes a more or
less compact white band in middle of body whorl, the upper part of which
appears above the suture in preceding whorls. Worn specimens with the latter
pattern may appear at first glance like latruncularia, but the larger number of
spiral striae easily distinguishes them. Protoconch white, brown patches begin
to appear on 2nd whorl (c. 4 mm. long.).
False Bay to East London.
Remarks. Protoconchs and juveniles from Still Bay in the Muir collection
(S. Afr. Mus.). Not taken by the Pieter Faure.
Mitra aerumnosa Melv.
Fis: 11(a)
1888. Melvill. 7. Conch., v, p. 282, pl. 2, fig. 12.
1903. Von Martens. D. Tiefsee Exp., vil, p. 31 (sumplex, non Dnkr.).
1919. Cooke. loc. cit., p. 416 (radula).
1925. Thiele. D. Tiefsee Exp., xvii, p. 185, pl. 32 (20), fig. 22 (simplex, non Dnkr.).
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 49.
Aperture subequal to spire. Protoconch 2 whorls, alt. 1 mm., diam.
0-75 mm., smooth. Postnatal whorls 6; spiral series of punctae 8-g on 2nd
whorl, 14-15 on body whorl, 16-18 additional series on base, becoming
punctate striae anteriorly; growth-lines feeble, but sometimes strong enough
to give a semicancellate appearance on the upper whorls. Columella pleats 4,
the lowermost one weak. Periostracum thin. 33 x II mm.
Buff with a paler median band on body whorl, sometimes only a series of
pale spots, sometimes also a series of pale dashes from the suture downwards,
periostracum yellowish, olivaceous, or castaneous.
Radula with c. 50 rows, central plate 4 times as wide as long, 8-cuspid,
lateral about 6 times as wide as long, 14-cuspid.
Fossil, late Tertiary; Saldanha Bay (Haughton).
Algoa Bay (Sowerby); Table Bay and Dassen Island (S. Afr. Mus.) ;
Natal (S. Afr. Mus.). Living: Oudekraal (west coast of Cape Peninsula) and
Langebaan (Saldanha Bay) (U.C.T.).
Remarks. The Natal specimen was received from Col. Bowker, but
probably not collected in Natal; Turton did not obtain it at Port Alfred; and
even the Algoa Bay record is open to doubt. lf
en nero Soar |
am
SENT mea
LSE Gye
Central and lateral radula plates of (a) Mitra aerumnosa Melv.; (6) M. litterata Lam.; (c) M.
(Dibaphus) bathybius n. sp.; (d) Vexillum sculptile (Rve.); (e) V. capense (Rve.); (f) Pusia patula
Rve.).
42 ANNALS OF THE SOUTH AFRICAN MUSEUM
nw See =
Thiele disagreed with von Martens’s identification of specimens from
34° 51'S. 19° 37’ E., 80 metres as simplex Dnkr.; he figured one but without
deciding its specific identity. The figure looks very like an aerumnosa.
A series in the Juritz collection (S. Afr. Mus.), probably from the west
coast of the Cape Peninsula, contains specimens from 5 mm. long upwards.
These show that the lowermost columella pleat does not develop until the
shell is 14-16 mm. in length (4 postnatal whorls), and even in adults always
remains feeble.
Mitra teretiuscula Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 185, pl. 32 (20), fig. 23.
Spire subequal to aperture. Protoconch 14 whorls, alt. and diam.
0°75 mm., smooth. Postnatal whorls 5, profile convex, suture deep but not
canaliculate; 4-5 (Thiele said a variable number) fine spiral striae, the one
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 43
below the suture usually a little more conspicuous than the others, none on
base below periphery; growth-lines arcuate. Columella pleats 4. 9°5 x 3-5 mm.
Yellowish with brown markings: an infrasutural and a median band, with
zigzag axial flames (Thiele); the present specimens show only the axial flames.
35° 16’ S. 22° 26’ E., 155 metres (Thiele) ; off Gape St. Blaize, 125 fathoms,
3 dead; off Cape Morgan, 45 fathoms, 2 dead (S. Afr. Mus. P.F. coll.).
Remarks. The rounded profile of the whorls, and the fewer spiral striae
distinguish this species from aerumnosa. Moreover the latter is a larger species
with only 3 whorls at 9 mm. length. Thiele’s figure shows the columella pleats
more oblique than in the present specimens.
In slightly worn specimens the uppermost stria (below the suture) persists
when the others have become untraceable.
Mutra (Papalaria) episcopalis Linn.
1833. Quoy & Gaimard. Voy. Astrolabe. Moll., pl. 45, figs. 1-7 (living animal).
1859. Chenu. Man. Conchyl., i, fig. 996.
1880. Von Martens. Mauritius G Seychellen, p. 249.
1919. Cooke. loc. cit., pp. 406, 408 (radula).
1935. Dautzenberg. Mem. Mus. R. Hist. Nat. Belg., H.S. 11, 17, p. 44 (references).
Radula with 71 rows, central plate 8—g-cuspid, lateral 20-22 cuspid, the
cusps diminishing to mere serrations at outer end (Cooke).
Distribution. Mozambique (Smith), Zanzibar, Mauritius, Madagascar,
Indo-Pacific.
S. Afr. Mus. has this well-known species from Mozambique.
Von Martens records that the living animal exudes a purple-brown fluid
which stains the hands reddish-brown, and smells like green walnuts.
Mitra (Papalaria) pontificalis Lam.
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 198.
1859. Chenu. Man. Conchyl., i, fig. 1030.
1880. Von Martens. Mauritius G Seychellen, p. 250.
1935. Dautzenberg. loc. cit., p. 53, pl. 2, figs. 8, g coloured (and var. confluens) (references).
Three specimens in the Ross-Frames collection (S. Afr. Mus.) labelled as
from Mozambique.
Distribution. Zanzibar, Mauritius, Madagascar, Indo-Pacific.
Mitra (Strigatella) limbifera Lam.
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 214.
1859. Chenu. Man. Conchyl., i, fig. 1003.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 44 (cartfa).
1919. Cooke. loc. cit., pp. 410, 411 (radula).
1935. Dautzenberg. loc. cit., p. 88 (aurantia var. |.) ( references).
1936. Peile. loc. cit., p. 142 (radula).
Aperture a little longer than spire. Postnatal whorls 8. Body whorl with
8 spiral grooves, varying in strength, often with fine spiral striae on the flat
44 ANNALS OF THE SOUTH AFRICAN MUSEUM
ridges, 15-18 additional grooves on base, becoming stronger anteriorly; crossed
by close-set slightly retractive growth-lines. Columella pleats 5, the lowermost
one usually feeble, sometimes with weak intermediary pleatlets. Periostracum
thin. 38 < 16 mm. (S. Afr. Mus.).
Upper whorls and upper half of body whorl yellowish-brown, base
darker chestnut brown, periostracum yellowish.
Radula with 86 rows, central plate 7-cuspid, lateral plate with 9-10 short
thick cusps, margin external to them bare (Cooke).
Natal (Krauss, Sowerby, S. Afr. Mus.) ; Durban (Sowerby, Smith, Cooke).
Distribution. Mauritius, Indo-Pacific (aurantia), Philippines (limbzfera).
Remarks. The spiral sculpture varies in strength, and one can speak of 8
spiral grooves or of 4 spiral flat ‘cords’ with lirae (carifa); the spiral grooves
are constant on the early whorls, variation occurring on the body whorl; in
one specimen the spiral sculpture on the body whorl shows only very faint lirae.
Mitra (Strigatella) litterata Lam.
Fig. 11(d)
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 220.
1859. Chenu. Man. Conchyl., i, fig. 1000.
1919. Cooke. loc. cit., pp. 410, 411, fig. 6 (radula).
1935. Dautzenberg. loc. cit., p. 103 (references).
1936. Peile. loc. cit., p. 142 (radula).
Aperture twice as long as spire. Postnatal whorls 6. Last whorl with 7
spiral rows of punctae (fewer on early whorls), 15-18 additional rows on base,
becoming grooves anteriorly. Columella pleats 4. Outer lip with rounded boss
in middle internally. Periostracum thin. 30 x 16 mm. (S. Afr. Mus.).
White with brown axial irregular zigzag or undulate markings, perio-
stracum yellowish.
Radula with c. 85 rows, central plate with 6 cusps, outermost cusp on
either side small, lateral plate wide with 8-10 cusps, outer third of margin
smooth. Anterior quarter of radula yellowish-brown, especially dark in front,
hinder three-quarters colourless.
Living: Durban and Kosi Bay (U.C.T.); Delagoa Bay (U.W.).
Distribution. Red Sea, Aden, Mauritius, Indo-Pacific.
Remarks. Cooke gave the number of cusps on the central plate of the
radula as 7, but his fig. 6 shows only 5; the outermost one on either side is
evidently too minute to show in the figure. Peile said the central plate may
have ‘one more cusp’ (?, 1.e. 8). The example here described, from Delagoa
Bay, has no median cusp.
Mitra (Strigatella) luctuosa A.Ad.
1844. Reeve. Conch. Icon., fig. 94 (polita).
1851. A. Adams. Proc. Zool. Soc. Lond., p. 133.
1880. Von Martens. Mauritius & Seychellen, p. 250 and p. 252, pl. 20, fig. 15 (polita Rve.).
1919. Cooke. loc. cit., pp. 410, 411 (radula).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 45
Aperture a little shorter than spire. Postnatal whorls 7. Last whorl with
7 spiral punctate grooves (fewer on early whorls), c. 12 additional grooves on
base, becoming stronger anteriorly. Columella pleats 4. Periostracum thin.
30 X 11 mm. (S. Afr. Mus.).
Chestnut brown, with a pale narrow spiral band a little distance below
the suture: between the 3rd and 4th rows of punctae (von Martens: on the
grd row).
Radula with 81 rows, central plate small, 7-cuspid, outermost cusp on
either side very small, lateral plate wide with 7—8 cusps and a long bare margin
external to them (Cooke).
Durban (Sowerby, Cooke). Farquhar Island (S. Afr. Mus.).
Mitra (Scabricola) texturata Lam.
1811. Lamarck. Ann. Mus. Paris, xvii, p. 213.
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 8.
1935. Dautzenberg. Mem. Mus. R. Hist. Nat. Belg., H.S. II, 17, p. 84, pl. 3, fig. 5 coloured
(references).
A specimen in S. Afr. Mus. collected by L. E. Kent on the Natal south
coast (between Durban and Port Shepstone) confirms Sowerby’s record from
Durban. It is slightly worn but retains the coloration (cf. Dautzenberg). The
species is therefore presumably an inhabitant of South African waters.
Distribution. East Indies, Philippines.
Mitra (Scabricola) crenifera Lam.
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 204.
Igit. Schepman. Siboga Exp. monogr., xlix, 1, p. 271, pl. 23, fig. 3 (radula).
1936. Peile. loc. cit., p. 143 (radula).
Aperture slightly longer than spire. Protoconch 2 (24) whorls, alt. and
diam. 0-75 mm., smooth. Postnatal whorls 7, profile slightly and evenly convex.
Axial ribs ¢. 20 on Ist and 2nd whorls, 24 on 3rd, increasing to c. 60 on 6th, and
very numerous on last whorl, especially on back of outer lip, arcuate, separated
by narrow grooves, extending across base; crossed by spiral lirae, 3 on 1st and
end whorls, 4 on 3rd and 4th, increasing to 1o on last whorl, some of them
with a subsidiary narrow lira, c. 14 additional lirae on base; sculpture cancel-
late, formed of small squares on most of the surface, but of axial oblongs on
last whorl, mostly smooth but slightly nodulose on base. Columella pleats 4.
42 X 14°5 mm., width of 3rd whorl 2-75-3 mm. (contrast circula, rufescens).
Cream or buff, with pale orange irregular spots below the suture, lower
half of whorl brown with curved retrorse flames projecting upwards, base with
a similar brown band with projections, columella glaze with orange margin.
Radula with c. 60 rows, central plate about twice as broad as long, with
2 strong cusps flanked by 2 smaller ones on either side, lateral plate 24 times
as broad as long, with 4 strong cusps on inner half, external half of margin bare
(Schepman, Peile).
46 ANNALS OF THE SOUTH AFRICAN MUSEUM
Durban (Sowerby); Delagoa Bay, fresh, presumably living, but no
specimen with animal (U.W.).
Off Umhlanga River (Natal), 22-26 fathoms, 1 dead, 18 mm.; off
Umkomaas (Natal), 40 fathoms, 1 broken, but with protoconch, 12 mm. (S.
Ate Mas.oP BE. 'coll.).
Distribution. Indian Seas, Philippine Islands, Fii, Ceylon, Nicobars,
Andamans, Mauritius.
Remarks. I am indebted to Mr. A. E. Salisbury for the identification of
this species.
The apical whorls increase in width more rapidly than in circula, rufescens,
acutilivrata.
Mitra ‘circulata Kien. var.’
1919. Cooke. loc. cit., p. 415 (radula).
Radula with c. 73 rows, central plate square, with 2 strong cusps flanked
on either side by 2 small obscure denticles; lateral plate transversely oblong,
with 4-5 cusps (Cooke).
Remarks. The radula described by Cooke from a Durban specimen
corresponds so closely with that of crenifera that it raises the suspicion that it
was extracted from a crenifera and not from one of the shells later described as
burnupiana (see p. 49).
Mitra flammea Q. & G.
1833. Quoy & Gaimard. Voy. Astrolabe Moll., p. 659, pl. 45 bis, figs. 23-5 (living animal).
1844. Reeve. Proc. Kool. Soc. Lond., p. 173 (interlirata).
1919. Cooke. loc. cit., p. 413, fig. 11 (radula), and p. 414 (interlirata) and footnote
(synonymy).
1925. Thiele. D. Tiefsee Exp., xvii, p. 185 (flammigera).
Remarks. Smith (1903. Proc. Mal. Soc., v, p. 366) records flammigera from
‘Kalk Bay (Burnup)’. Some error seems to have occurred, because it is very
unlikely that this Indo-Pacific species extends (certainly not living) as far west
as Kalk Bay on the east side of the Cape Peninsula. Probably the collector’s
name (Burnup) should refer to the locality Durban, not to Kalk Bay.
Mitra rufescens A.Ad.
Fig. 12(d)
1851. A. Adams. Proc. Zool. Soc. Lond., p. 137.
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 9 (? if distinct from circulata [= circula]).
Aperture subequal to spire. Protoconch ? (corroded). Postnatal whorls
7. Spiral lirae 3 on all whorls, but on 6th and 7th a 4th low down and almost
concealed in the suture, a weaker intermediary lira between each pair on
5th—7th whorls, and 1-2 striae on last whorl (or the lowermost intermediary
—"
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 47
lira may be duplicated) ; lirae cut by 15-16 narrow axial grooves on 2nd whorl,
increasing to c. 24 on 5th, and c¢. 50 on last whorl; sculpture cancellate with
squares on early whorls and axial oblongs on later whorls, but spiral lirae
predominant. Columella pleats 3, sometimes a feeble 4th. 26 x 9 mm.,
width of 3rd whorl 1-5-1-75 mm.
Buff, more or less rufescent, possibly due to staining (iron stanchions, etc.,
in Durban harbour).
Durban (Sowerby; also S. Afr. Mus. ex coll. Ross-Frames, fresh,
presumably living).
(iste eeee puntata as see
itt psnnasaneeen csassimens sees 3 .
Rar ett Lessee esees==
Ave ererrrre eae SSeeseees:
poet err ler os ee a eaees
Lee
SSS — SS eee=
ATI
isi
teense
i
Bre? 12:
One whorl (semi-diagrammatic) of (a) Mitra circula Kien.; (6) M. rufescens A. Ad.; (c) M. acuti-
lirata Sow.; (e) M. foveolata Dnkr.; (d) protoconch and first two postnatal whorls of M. acutilirata
Sow.
Mitra acutilirata Sow.
Fig. 12(c), (d)
1874. Sowerby. Thes. Conch., iv, sp. 129, pl. 15, figs. 273, 274.
1936. Peile. loc. cit., p. 142 (radula).
Aperture subequal to spire. Protoconch 4 whorls, somewhat lopsided, alt.
0:8, diam 0-6 mm., smooth. Postnatal whorls 8. Spiral lirae 3 at start of Ist
whorl, 4 on each of gnd—6th whorls, 5 on 7th and 8th whorls, 7-8 additional
lirae on base; these main lirae rather sharply subcarinate, between them 1-3
striae and sometimes a weaker intermediary subcarinate lira on last whorl;
lirae cut by 13-14 narrow axial grooves on 2nd whorl, increasing to ¢. 22 on
5th and 45-50 on last whorl; sculpture cancellate with small squares and axial
oblongs. Columella pleats 4, the lowermost one feeble. 26 x 9 mm., width of
3rd whorl 1-5-1-75 mm.
48 ANNALS OF THE SOUTH AFRICAN MUSEUM
Dull greyish, a brownish band around middle of body whorl, with some
irregular flame-like expansions above (cf. crenifera), the main lirae with small
brown spots on their crests.
Radula of the Mitra (s.s.) form, central plate with c. 10 cusps, lateral plate
very wide, with cusps throughout its width (Peile).
Delagoa Bay, 1 juv. (S. Afr. Mus. coll. K.H.B. 1912); same locality, 3
(U.W.).
Distribution. Farquhar Island (S. Afr. Mus. coll. E. L. Layard, on board
H.M.S. Castor, 1856); Mauritius, Indo-Pacific.
Remarks. 1 am indebted to Mr. A. E. Salisbury for the identification of
this species.
A Philippine specimen 37 X 13 mm. (S. Afr. Mus. ex coll. Ross-Frames)
named scabriuscula Linn. (by Sowerby & Fulton) agrees with the above
description except there are 5 main lirae on 5th whorl, 6-7 on 6th, 7-8 on 7th
whorl. The increase of the intermediary lirae to become main lirae is scarcely
of specific importance; but I am not able to decide the validity of Sowerby’s
species, and accept Mr. Salisbury’s identification. Neither scabriuscula nor
acutilirata has hitherto been recorded from South Africa.
Three dead juveniles from off O’Neil Peak (Zululand), 90 fathoms (S. Afr.
Mus. no. A8811. P.F. coll.) may be this species, but the axial grooves are more
numerous than in the other specimens at hand.
Mitra (Cancilla) circula Kien.
Fig. 12(a)
1839. Kiener. Spec. Cog. Mitra, p. 21.
1919. Cooke. loc. cit., p. 417 (radula).
1921. id. Proc. Mal. Soc., xiv, p. 114, fig. 2 (burnupiana).
1936. Peile. loc. cit., p. 143, fig. 9 (radula).
Aperture subequal to spire. Protoconch 34 whorls, diam. 0-6, alt. 0-75 mm.
Postnatal whorls 8. Spiral lirae 3 on each whorl, from 4th or 5th onwards a
4th lira low down and partly concealed in the suture, 1-3 striae and sometimes
a subsidiary lira above Ist lira, 1-2 striae between each pair of main lirae and
sometimes 2 lirae between 3rd and 4th lirae, 7 (8) additional lirae on base, with
intermediaries; lirae cut or nicked by numerous close-set axial striae, 15-16 on
1st whorl, 19-20 on 2nd, increasing to at least 60 on 7th and more on 8th whorl ;
sculpture cancellate with small squares and axial oblongs, lirae predominant.
Columella pleats 3 with an obscure 4th. 31:5 x 10 mm., width of 3rd whorl
1°75-2 mm.
White or buff, lower half of whorls faintly orange.
Radula with arcuate central plate, lateral plate transversely elongate,
both plates with numerous denticles or cusps (Cooke, Peile).
Inner harbour, Durban (S. Afr. Mus. coll. Ross-Frames); Cape Natal,
54 fathoms, 1 broken apex (S. Afr. Mus. P.F. coll.); Delagoa Bay (U.W.).
3
CONTRIBUTIONS TO KNOWLEDGE OF 58.A. MARINE MOLLUSCA 49
Remarks. Except for the development of more numerous axial striae, there
seems little difference between the specimens here referred to circula and those
referred to rufescens.
The status of the name burnupiana depends on the shell alone, because the
_ description was based on the shell and was accompanied by a figure (albeit
diagrammatic). It is here treated as a synonym.
The distinctions (Cooke, 1921) between crcula and burnupiana, both found
together in Durban Bay, are not very convincing. Plump and slender indivi-
duals occur in many Gastropods. The incidence of the line joining the
columella pleats (see Burnup’s diagram in Cooke, 1921) depends on the angle
from which the shell is viewed, and in any case may be vitiated by a very slight
curvature in the columella, or the variable development of the pleats.
It seems most unlikely that there should be two species living together
which are scarcely distinguishable conchologically, but which have quite
dissimilar radulae (see p. 45, crenifera).
Mitra foveolata Dnkr.
Fig. 12(e)
1863. Dunker. WNovitat. Conch., p. 46, pl. 15, figs. 5, 6.
Fusiform. Spire 14 in aperture. Protoconch ? 3 whorls (broken). Post-
natal whorls 8. Spiral lirae 3 on each whorl, from 4th onwards a 4th lira low
down and. partly concealed in the suture, an impressed stria (sometimes two
striae) between each pair of lirae, between 3rd and 4th lirae a weak inter-
mediary lira, c. 10 additional main lirae on base, sometimes with intermediaries
between the upper 6 or 7 lirae; lirae cariniform, uninterrupted; between the
lirae close-set axial striae, often subfoveolate underneath each lira. Suture
slightly canaliculate. Columella pleats 4, the 4th feeble, but sometimes a
feebler 5th pleat. 25 < 9-10 mm.
Pale buff (sometimes stained orange-rufous) with irregular orange-brown
axial streaks or flames.
Durban (S. Afr. Mus. ex coll. Ross-Frames); Delagoa Bay (S. Afr. Mus.
coll. K.H.B., 1912).
Remarks. The identification of this species also I owe to Mr. Salisbury.
The continuous carinate lirae, not cut by the axial striae, distinguishes
the species from the others here recorded as acuitzlirata, rufescens, circula. Watson’s
figure of rufescens, however (1886. Challenger Rep., xv, pl. 14, fig. 5), shows a
similar sculpture.
Von Martens in 1880 (Mauritius G Seychellen, p. 251) gave this species as a
synonym of inferlirata Rve. 1844, and in 1903 (D. Tiefsee Exp., vii, p. 106) as a
synonym of flammigera Rve.
50 ANNALS OF THE SOUTH AFRICAN MUSEUM
Mitra (Swainsonia) ocellata (Swains.)
1831. Swainson. ool. Illustr. (2), ii, pl. 54 (Mitrella o.).
1854. H. & A. Adams. Gen. Rec. Moll., i, p. 180 (Swainsonia o.).
Fusiform. Aperture slightly longer than spire. Protoconch broken. Post-
natal whorls 7. Spiral punctate striae 3 on 1st whorl, 4 on 2nd, increasing to 5
on last whorl, 9 additional striae on base, not becoming stronger anteriorly;
fine close-set axial growth-lines. Columella pleats 5; a narrow parietal callus
from top of aperture to Ist pleat; outer lip slightly thickened. 27 X 9:5 mm.
Pale fawn with irregular white marks which are flame-like on upper
whorls, then breaking up into arrows and triangles, resembling on the body
whorl the pattern of a ‘textile’ cone.
Natal (S. Afr. Mus. received from a private collector, the locality therefore
not certain; also a worn specimen from the coast between Durban and Port
Shepstone collected by L. Kent).
Distribution. East coast of Africa, Singapore.
Remarks. The shell is apparently not much worn although highly polished.
The body whorl has three times suffered injury and been repaired; this may
account for the 5th columella pleat (though 5 pleats is usual in Swaznsonia) ; at
two of the breaks the outer lip shows a slight narrow thickening before the
continuation of growth.
M. fissurata Lam., of which ocellata has been considered a variety (Tryon.
Man. Conch.) occurs at Mauritius and Seychelles (1880. Von Martens.
Maunt. G Seych., p. 249); ocellata is dintinguished by the punctate-striate
sculpturing (A. E. Salisbury in litt. 1957).
Mitra (Callithea) subulata (Lam.)
Fig. 13(a)
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 211.
1859. Chenu. Man. Conchyl., i, fig. 1019.
Slender, fusiform, profile of spire straight. 6 whorls preserved (proto-
conch and ? 2 whorls missing). First 4 preserved whorls each with 4 spiral
striae; 5th whorl with 6, 6th with 2 plus 4 (or 5) indistinct striae. On the 4th,
5th and 6th whorls the uppermost stria is better developed than the others,
forming a Terebra-like groove. Indications of numerous low rounded, close-set
axial ribs on last 2 or 3 whorls, more distinct above the spiral groove than below
it, and making the suture undulate or crenulate. 16 or 17 additional striae on
base, becoming stronger anteriorly. Columella pleats 4. 25:5 x 8 mm.,
aperture 12 mm.
Off O’Neil Peak (Zululand), 90 fathoms, one dead and partly corroded
(S. Afr. Mus. no. A8763. P.F. coll.).
Remarks. Somewhat similar in shape to picta and aerumnosa, but the
distinctive feature is the Terebra-like spiral groove.
CONTRIBUTIONS TO KNOWLEDGE OF 5$.A. MARINE MOLLUSCA Bf
Mitra (Dibaphus) bathybius n. sp.
Figs. 11(c), 13(d)
Protoconch 2 whorls, alt. 1, diam. of 1st whorl 1, of 2nd whorl 1-5 mm.,
smooth. Postnatal whorls 4 (preserved), with cancellate sculpture; 1st whorl
at beginning with close-set axial riblets (some of which may belong to the
protoconch), later with in addition 4 spiral striae; arcuate axial ribs c. 20. on
ist whorl, 20 on 2nd, 23 on 3rd, and 25 on 4th whorl, distinct and sharp on
first three whorls, becoming feeble on 4th and indistinct towards outer lip;
spiral striae 5 on 2nd, 6 on 3rd, 7 on 4th whorl, crossing the ribs, broader and
deeper (almost grooves) on upper part of whorl than on lower part; 10
additional grooves on base (c. 20 including those on rostrum). No columella
pleats. No operculum.
apet’s
=t.
we ot SS ak
rast
q
Wig
I
\\i
\
Ay
\\
CCK
ti \ \
\
i
Pics 43-
(a) Mitra (Callithea) subulata Lam. (6) Mitra (Dibaphus)
bathybius n. sp.
Radula with 60 rows, central plate with 5 cusps, a tiny cusp or denticle
at side not traceable, lateral plate with 10 cusps, a small one at inner end,
followed by 3 large ones, the other 6 farther apart and decreasing in size.
Off Cape Natal, 440 fathoms, 1 alive and one fragment (S. Afr. Mus.
A8627. P.F. coll.).
Remarks. Very fortunately this shell, although preserved dry for 57 years
in a bottom-sample, contained a well-preserved animal; presumably the
sample had originally been preserved in formalin. Owing to the likeness of
the shell to Tomlin’s description and figure of Charitodoron pasithea (p. 146) it
was unexpected to find the animal possessed a Mitrid radula.
52 ANNALS OF THE SOUTH AFRICAN MUSEUM
The radula resembles that of M. (Dibaphus) edentula (see: Cooke 1919);
the shell, however, is very different in appearance from that of edentula.
It seems safer not to include the genus Charitodoron in the Mitridae, but to
wait until the radula of an undoubted species of this genus has been obtained.
Gen. VExILLUM Bolten-Réding
(syn: Turricula Klein)
Fusiform or turriform. Radula with arcuate, multicuspid central plate,
and simple falcate lateral plate.
Vexillum alauda (Sow.)
1874. Sowerby. Thes. Conch., iv, pl. 361, figs. 134, 135.
1903. Smith. Proc. Mal. Soc., v, p. 367.
1919. Cooke. loc. cit., p. 418 (radula).
Narrow fusiform. Aperture a little shorter than spire. Protoconch ?
Postnatal whorls 9. Axial ribs 14-15 on Ist whorl, 16-17 on 2nd, from the
5th or 6th whorl decreasing to 13 on the last whorl, thin, straight, slightly
shouldered on body whorl, descending to base, not crossed by spiral striae;
spiral striae 3-4 on Ist and 2nd whorls, increasing to 6—7 on last whorl, some
of them double, ¢. 15 additional ones on base becoming grooves anteriorly;
well impressed on early whorls, but becoming feebly so on later whorls.
Columella pleats 4. Outer lip feebly plicate internally. 40 (protoconch and
3 whorls missing) X 13 mm.
Fawn or grey, upper half of whorls with a brown band, bordered below
by a narrow line, a second brown band at bottom of whorl partly concealed in
suture, but clear on body whorl as far as outer lip, base anteriorly brownish;
half-grown specimens chestnut brown, with or without the pale yellowish line;
some specimens bluish-grey with two series of orange-brown spots on the ribs.
Radula, central plate with 16-17 denticles (Cooke).
Durban (Smith; and S. Afr. Mus); Inhambane, Portuguese East Africa
COR Fd tyne
Distribution. Mauritius.
Remarks. The specimens from the Morrumbene estuary, Inhambane, are
discoloured and corroded, but one retains the two series of brown spots on the
ribs.
Vexillum exasperatum (Gmelin)
1790. Gmelin in Linn. Syst. Nat., ed. 13, i, p. 3453 (Voluta e.).
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 219 (arenosa).
1911. Schepman. Szboga Exp. monogr., xlix, 1, p. 287 (torulosa as synonym, arenosa as variety).
1919. Cooke. loc. cit., p. 418, fig. 17 (radula), and p. 418 (arenosa).
1935- Dautzenberg. loc. cit., p. 148 (with vars. torulosa and arenosa) (references).
1936. Peile. loc. cit., p. 143, fig. 10 (radula).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 53
Cooke figured the central plate of the radula of exasperatum with 15-16
denticles; Peile’s figure shows 19 cusps, the median one the largest; and Cooke
gave 20-22 denticles for arenosum.
Durban (Sowerby); Delagoa Bay (U.W.); Mozambique Island (S. Afr.
Mus. coll. K.H.B. 1912).
Distribution. Red Sea, East Africa, Indo-Pacific.
Remarks. Two broken specimens were identified by Tomlin as arenosa.
They do not exactly correspond with specimens from New Caledonia identified
as this species by Sowerby & Fulton.
Vexillum torulosum (Lam..)
1811. Lamarck. Ann. Mus. H. N. Paris, xvii, p. 216.
1859. Chenu. Man. Conchyl., i, fig. 1014.
Turreted. Aperture a little shorter than spire. Protoconch ? Postnatal
whorls 6 (? 7), profile angularly shouldered. Axial ribs 15 on the apical two
whorls (probably the 2nd and 3rd), decreasing to 11 on last (6th or 7th) as
wide as intervals on early whorls, much narrower on later whorls, extending
across base; crossed by 2 spiral striae on early whorls, increasing to 4 on 4th
(? 5th) and 12 (5 above, 7 below shoulder) on last whorl, 10 additional striae
on base before the costate rostrum which has 3—4 lirae; growth-lines increasing
in strength and forming a cancellate sculpture on later whorls. Columella
pleats 4. 22 (apex missing) X II-5 mm.
Uniform dull fawn.
Delagoa Bay, one dead but fresh (U.W.).
Remarks. The specimen corresponds with Chenu’s figure, and is quite
distinct from the specimens identified by Tomlin as arenosum.
Vexillum daedalum (Rve.)
Fig. 14(a)
1845. Reeve. Proc. Zool. Soc. Lond., p. 54; and Conch. Icon., Mitra, pl. 34, fig. 281.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 227.
1911. Schepman. Siboga Exp. monogr., xlix, 1, p. 285.
Aperture 14 in spire. Protoconch broken. Postnatal whorls 8, sutures
deep but not canaliculate. Axial ribs 11 on ist whorl, increasing to 15-17 0n
last whorl, moderately sharp, extending across base, intervals bluntly V-shaped;
spiral grooves in the intervals 5 on 2nd whorl, increasing to 7 on last whorl,
8-9 additional grooves on base, deeply impressed in the intervals but not
crossing the ribs except on lower part of base where they cause the ribs to be
slightly nodulose. Columella pleats 4; a nodular parietal callus at top of
aperture. Outer lip plicate within. 19g X 7 mm.
Pale orange-brown (probably faded), a white band at top of whorl (below
the suture) and another at bottom of whorl (above suture), and one or two on
base of body whorl.
54 ANNALS OF THE SOUTH AFRICAN MUSEUM
Off Scottburgh (Natal), 92 fathoms, dead shells (Sowerby).
Off Umhlangakulu River (Natal), 50 fathoms, 2 dead (seen by Sowerby) ;
off O’Neil Peak (Zululand), 55 fathoms, 2 dead (S. Afr. Mus. P.F. coll.).
Distribution. Philippine Islands, East Indies.
Remarks. Having no authentic daedalum for comparison, I accept Sowerby’s
identification of two Natal examples (S. Afr. Mus. no. A3482), returned by
him but from a locality different from that which he recorded (1903). He did
not state the number of P.F. specimens received by him. One wonders whether
perhaps the two specimens included in a collection ‘from South Africa’ in the
United States National Museum, and described by Bartsch as helena, were P.F.
specimens (see p. 55).
al b Cc d
Fic. 14.
Sculpture (semi-diagrammatic) of (a) Vexillum daedalum (Rve.); (b) V. sculptile (Rve.); (d) V. dis-
coloria (Rve.); (c) protoconch and first four postnatal whorls of Vexillum sp. (S. Afr. Mus.
no. A8813).
An axial stria is sometimes present at the bottom of the V-shaped intervals;
and the V is not always symmetrical, the bottom being nearer to the following
rib than to the preceding rib. cf. discoloria.
Vexillum sculptile (Rve.)
Figs. 11(d), 14(b)
1845. Reeve. Proc. Zool. Soc. Lond., p. 55; and Conch. Icon., Mitra, pl. 35, fig. 290.
1874. Sowerby. Thes. Conch., iv, Mitra, p. 34, pl. 26, fig. 596.
1886. Watson. Challenger Rep., xv, p. 251.
1911. Schepman. Siboga Exp. monogr., xlix, 1, p. 286.
Seven specimens from Delagoa Bay differ in the following respects.
Aperture 14 in spire. Postnatal whorls 8-9. Axial ribs 15-16 on 1st whorl,
increasing to 26-30 on last whorl, not very sharp, extending across base,
intervals more or less V-shaped, narrower than in daedalum; spiral grooves in
the intervals 4 on 2nd whorl, increasing to 5 (6) on last whorl, 8-9 additional
CONTRIBUTIONS TO KNOWLEDGE OF 58.A. MARINE MOLLUSCA 55
grooves on base, deeply impressed in the intervals (in the largest specimen,
which has 30 ribs, they are more like pits than grooves), not crossing the ribs
except on lower part of base where the ribs are slightly nodulose; the and
groove from the top of whorl is the deepest, usually cutting the ribs and forming,
especially on the early whorls, a Yerebra-like continuous spiral groove. Colu-
mella pleats 4; a nodular parietal callus at top of aperture. Outer lip plicate
Within. 21 X 7 mm.
Upper half of whorls more or less brown, lower half cream, base brown.
Radula (only one available) with 55 rows, central plate concave in front,
with 13 cusps between the acute postero-lateral angles, lateral plate falcate.
Two of the cusps on the central plate often united to form a single apically
bifid cusp, the fusion not occurring on every plate, and not in the same position.
Inhaca Island, Delagoa Bay, 2 living and 5 dead (S. Afr. Mus. coll.
K.H.B. 1912).
Distribution. Kast Indies, Philippine Islands.
Remarks. Bartsch (1915. Bull. U.S. Nat. Mus., 91, p. 43) described Mitra
helena based. on two specimens from “South Africa’, originally sold by Sowerby
& Fulton as daedalum. Bartsch said helena (15 * 6-6 mm.) was smaller (how
much smaller?) than the Philippine daedalum. The description gave the
numbers of axial ribs as 16 on Ist whorl, 20 on 2nd, and 16 [szc] on the last; if
16 is a typ. err. for 26, the numbers correspond with those of the Delagoa Bay
specimens better than with those of the specimens here recorded as daedalum.
Vexillum sp.
Fig. 14(c)
A single specimen in fresh condition taken off Cape Natal (Durban), 54
fathoms (S. Afr. Mus. no. A8813. P.F. coll.) is closely allied to daedalum, and
may be referable to sculptile, but is much more slender.
Narrow fusiform. Aperture 1$ in spire. Protoconch 3% whorls, slightly
lopsided, alt. 0-75, diam. 0-5 mm., smooth. Postnatal whorls 74. Sutures deep,
but not canaliculate. Axial ribs 11-12 on 1st whorl, increasing to c. 22 on 4th
and ¢. 33 on last whorl, subequal in width to the intervals, extending across
base; spiral striae 3-4 on 1st whorl, from 2nd whorl onwards the uppermost
stria becomes groove-like, more deeply impressed, and cuts the ribs, followed
by 4 striae, forming rather deep pits in the intervals but scarcely cutting the
ribs; from 4th whorl onwards an additional feeble stria is developed between
the suture and the deeply impressed stria; 7 additional striae on base. Colu-
mella pleats 4; a feeble nodular parietal callus at top of aperture. Outer lip
plicate within. 15 X 4°5 mm. (probably not quite fully grown).
White, a very faint trace of a yellowish band below suture and another
on lower part of base of body whorl.
56 ANNALS OF THE SOUTH AFRICAN MUSEUM
Vexillum discoloria (Rve.)
Fig. 14(d)
1845. Reeve. Proc. Zool. Soc. Lond., p. 46; and Conch. Icon., Mitra, pl. 29, fig. 230.
1880. Von Martens. Mauritius G Seychellen, p. 255 (discolor [sic]).
Two P.F. specimens were identified by Sowerby as this species, but the
record was not published.
Fusiform; aperture 14-14 in spire. Protoconch broken. Postnatal whorls
7. Sutures deep but not canaliculate. Axial ribs 11 on 1st whorl, increasing to
15-16 on last whorl, rather sharp, much narrower than the more or less
V-shaped intervals, extending across base; a fine axial stria along the bottom of
the intervals, which is sometimes asymmetrical (cf. daedalum); spiral striae 5
on 2nd whorl, 7 on last whorl, c. 14 additional striae on base, strongly marked
in the intervals, not cutting the ribs except on base where the intersected ribs
are slightly nodulose; on 1st—3rd whorls the stria next the suture is the deepest,
at 4th whorl a weaker stria is interpolated, and on 5th and 6th whorl 2 striae;
thus the 1st, then the 2nd, and ultimately the grd stria is the strongest and most
deeply impressed, forming a Terebra-like spiral groove, especially noticeable on
the early whorls. Columella pleats 3, with an obscure 4th anteriorly. Outer
lip plicate within. 12°5 * 4°5 mm.
Pale brown, slightly darker in the intervals, with a white band around
middle of whorl, and two other bands on the base.
Off Umvoti River (Natal), 27 fathoms, 2 dead but fresh (S. Afr. Mus.
PF. coll.)
Distribution. Mauritius, Réunion (von Martens).
Remarks. Reeve’s description (I have not seen his figure) does not quite
fit the present specimens except as regards the colour: ‘fasciis roseis et albis
alternata, fasciis roseis inter costas ustulato-nigricantibus’.
Vexillum capense (Rve.)
Fig. 11(e)
1845. Reeve. Conch. Icon., Mitra, pl. 33, fig. 268.
1903. Von Martens. D. Tiefsee Exp., vii, p. 53 (listed) (Turricula (Pusia) c.).
1910. Schwarz. Tr. Geol. Soc. S. Afr., xii, p. 115.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 44, pl. 21, fig. 4 (ima).
1932. Turton. Mar. Sh. Port Alfred, p. 47, pl. 10, no. 354; and p. 47 (ima).
1932. id. ibid., p. 48, pl. xi, no. 356 (albanyana).
1932. id. ibid., p. 48, pl. xi, no. 357 (hera).
Aperture a little shorter than spire. Protoconch 14 whorls, alt. 0-5, diam.
o-6-0'7 mm., smooth. Postnatal whorls 5. Axial ribs 11-12 on 1st and and
whorls, 14-15 on body whorl, narrower than the intervals (in fresh specimens),
extending across base (in fresh specimens) ; spiral striae in the intervals 7 (8-9),
not crossing the ribs, often not conspicuous, 11-12 additional striae on base,
s
4
ae Se igde: bade
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 57
but usually only the 5-6 stronger anterior ones conspicuous. Columella pleats
4. 11°5 X 5 mm. (Turton: 15 mm. long.).
Pale fawn, with a darker orange-brown band, sometimes mottled, above
a white peripheral band, base orange-brown more or less spotted with white;
the white band is usually suffused in the middle so that an upper and a lower
band of pure white is formed; the axial grooves also are often suffused, which
produces a ‘necklace’ effect of white spots on the ribs, and the upper necklace
is bordered below by a narrow brown line (cf. Bartsch’s figure of ima).
Radula, one specimen examined but presumably incomplete because only
20 rows were obtained, central plate strongly arcuate in front, with 13 cusps
between the acute postero-lateral angles, lateral plate falcate.
Fossil: Pleistocene, Port Elizabeth (Schwarz).
Living: False Bay (U.C.T.).
Dead: Kalk Bay (False Bay) to Port Alfred (auct. et S. Afr. Mus.); Ton-
gaat (30 miles north of Durban) (S. Afr. Mus.); Still Bay (U.C.T.).
Off Nanquas Peak (eastern part of Algoa Bay), 63 fathoms, 3 dead; off
Cove Rock (East London), 22 fathoms, 1 juv. (S. Afr. Mus. P.F. coll.).
Remarks. Although Dunker is quoted as author by Krauss, Sowerby
(1892), Bartsch, and Turton, the species should be credited to Reeve (see
Sherborn. Index Anim.).
Neither Bartsch nor Turton seems to have realized that ima is merely a
worn specimen of capensis. Among a large number of beach-worn specimens
two or three can usually be found which have reached the right stage of
abrasion. The same applies to albanyana and hera.
Under capensis Turton (p. 47) mentions rufocincta Adams 1851 as a ‘variety
(or synonym)’; see ewzonata (p. 38).
Sometimes only 4 or 5 well-spaced spiral! striae are visible in the intervals,
e.g. the 3 P.F. specimens, and one out of about 50 specimens from Still Bay
(S. Afr. Mus.). A similar reduction in the number of spiral striae occurs in a
specimen of eucosmia Turton = kowieensis (p. 39).
Gen. Pusta Swainson
Fusiform. Radula with arcuate, tricuspid central plate, and simple falcate
lateral plate.
Pusia patula (Rve.)
Fig. 11( f)
? 1840. Kiister in Mart. Chemn. Syst. Conch. Cab., p. 108, pl. 17a, figs. 4-6 (pruinosa) (fide von
Martens, 1903).
21845. Reeve. Proc. Zool. Soc. Lond., p. 49; (M. pica) and Conch. Icon., Mitra, pl. 31, fig. 247.
1845. id. ibid., p. 61; and ibid., pl- 39, fig. 333 (M. patula).
1846. Dunker. eitschr. Mal., iii, p. 111 (simplex).
1848. Krauss. Stidafrik. Moll., p. 125, pl. 6, fig. 20 (simplex).
1889. Sowerby. 7. Conch., vi, p. 8, pl. 1, fig. 11 (merula).
1903. Smith. Proc. Mal. Soc., v, p. 366 (pica and patula).
58 ANNALS OF THE SOUTH AFRICAN MUSEUM
? 1903. Von Martens. D. Tiefsee Exp., vii, p. 31 (simplex, ? non Dnkr.).
1906. Smith. Ann. Natal Mus., i, p. 33 (simplex; removes the Tasmanian cinnamomea from the
synonymy).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 45 (merula, simplex and patula).
1916. Sowerby. Ann. Mag. Nat. Hist. (8), xviii, p. 491, text-fig. (fidis).
1922. Peile. Proc. Mal. Soc., xv, p. 94, fig. 4 (radula) (merula).
? 1925. Thiele. D. Tiefsee Exp., xvii, p. 185, pl. 32 (20), fig. 22 (? simplex, disagrees with
von Martens 1903).
1925. id. ibid., p. 186, pl. 32 (20), fig. 24 (hottentota).
1932. Turton. Mar. Sh. Pt. Alfred, p. 45 (merula); p. 45, pl. 10, no 337 (lurida); p. 45, pl. 10,
no. 338 (fidis); p. 46 (patula, pica and simplex); p. 46, pl. 10, no. 340 (simplex var.
alfredensis).
1938. Peile. Proc. Mal. Soc., xxiii, p. 100, fig. 37 (radula) (patula).
Ovate-fusiform. Aperture about 14 times the spire. Protoconch 2 whorls,
alt. and diam. 0-5 mm., smooth. Postnatal whorls 6-7. First whorl at start
with some fine axial pliculae, following whorls smooth, feebly or distinctly
ribbed, when present 12-14 ribs on 4th and 15-18 on last whorl; fine spiral
striae 3 on Ist whorl, increasing to 5 or 6 on upper part of whorls, minutely
crinkly (due to growth-lines) if whorls smooth, visible only in the intervals
when the whorls are ribbed; usually no striae visible on periphery, but 4-5
may appear on lower half of whorls; in fresh examples the striae probably
cross the ribs; on base 15-18 additional striae, usually visible only on lower
part where they separate c. to lirae. Columella pleats 4. 17-5 (apex worn)
x 6-5 mm.
Various shades of brown, sometimes very dark, almost black, often with
a white mark on outer side of outer lip, or with a white interrupted spiral band
in middle of whorl; some specimens mottled.
Radula with c. 70 (Peile: 64-67) rows, central plate arcuate, tricuspid,
lateral plate falciform. The shape of the lateral plate seems to vary; my
examples are intermediate between Peile’s figures of merula (1922) and patula
(1938).
From Durban along the south coast and around the Cape Peninsula up
to Port Nolloth on west coast.
35° 16'S. 22° 26’ E., 155 metres (Thiele: hottentota).
Remarks. M. pruinosa is listed above on the authority of von Martens (1903).
M. pica was described from an unknown habitat, but listed as South African
by Smith (1903), and Turton, though the latter did not claim to have found
any at Port Alfred. I have seen no specimens with a ‘jagged white band’
encircling the black whorls ‘next the sutures’ (Reeve, italics mine). A white band
is frequently present in the middle (or slightly above) of the whorl, and may be
almost continuous, or divided into a series of spots, or it may appear as a
white mark only on the outer lip.
Peile (1922) remarks that though ebenus (to which Sowerby likened his
merula) has a smooth shell, its var. savignyi is costulate; merula is ‘slightly costu-
late’. This lends support to the present contention that only one species is
found in South Africa.
4
y
f
;
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 59
The references to von Martens and Thiele are included with queries.
Thiele disagreed with von Martens’s identification, but left the specimens sub
judice; his figure strongly suggests that the dead and bleached Valdivia specimens
were really aerumnosa (q.v. p. 41).
Turton’s simplex var. alfredensis appears to be merely a redescription of
Reeve’s paiula: ‘cinerea, fusco hic illic variegata et nebulata’ (Reeve), ‘a light
mottled variety showing various colours’ (Turton).
Peile’s specimens of merula (Durban) and patula (East London) were both
supplied by Tomlin, who presumably favoured the recognition of two species.
Their radulae scarcely differ sufficiently to indicate two species; especially as
my examples are intermediate in shape.
I consider therefore that only one species, common around the whole
South African coast from Durban to Port Nolloth, should be recognized; but
I am not in a position to decide whether fica, having page precedence, should
replace patula, or whether the earlier pruinosa should be accepted.
The Pieter Faure obtained no examples except one very slender shell off
Cape Morgan, 47 fathoms (S. Afr. Mus. no. A8812. P.F. coll.). Pale brown.
Four whorls, 8 x 3 mm. (a normal 4-whorled example is 7-5 X 3:3-3°5 mm.).
13 ribs on last whorl; 13-14 spiral striae on last whorl, those nearest the suture
closer together than the others, as in normal examples.
Gen. ImBRICARIA Schumacher
Biconical. Radula as in typical Mitra.
Imbricaria carbonacea Hinds
1844. Hinds. Zool. Voy. Sulphur., p. 41, pl. xi, figs. 9, 10.
1910. Dautzenberg. Act. Soc. Linn. Bordeaux, p. 47.
1912. id. Ann. Inst. oceanogr., V, fasc. 3, p. 28.
Smooth, polished, with obscure grooves on base; columella pleats 5, the
lower ones obscure. Size not stated: Hinds’s figure 22 XxX 11 mm. Black,
aperture internally pale.
Baie de Rufisque (Dakar, West Africa), 18-20 metres (Dautzenberg 1910) ;
Mossamedes, 15-20 metres (Dautzenberg 1912).
Remarks. Described by Hinds from deep water on the Agulhas Bank, but
it has not since been recorded from South African waters. Probably some
error occurred in labelling Hinds’s material, as Belcher (in command of the
Sulphur) is known to have relied upon his memory.
Fam. OLIVIDAE
1929. Thiele. Handbuch, i, p. 330.
On account of its radula Thiele included Melapium in this family, although
the shape of the shell is very different from that of any of the other genera.
60 ANNALS OF THE SOUTH AFRICAN MUSEUM
Thiele (p. 332) created the genus <emiropsis for Eburna (i.e. Babylonia)
papillaris Sow., stating that the systematic position was doubtful in the absence
of knowledge of the operculum and animal. He evidently overlooked Sowerby’s
figure (1902. Mar. Invest. S. Afr., 11, pl. 2, fig. 3); and moreover gave no con-
chological reasons for instituting a new genus or for including it in the Olividae.
In fact this species is a true Babylonia (p. 147).
Gen. OLIva Brug.
The Pieter Faure obtained one very worn ispidula Linn. from Morewood
Cove (between Umhloti and Umvoti, Natal), 27 fathoms; one living dactyliola
Duclos 1835 from off Tugela River, 14 fathoms; and one dead dactyliola from
off Umkomaas, 13 fathoms (the first two identified by Sowerby 3rd).
Braga (1952. Anais 7. Invest. Ultramar., vii, 3, p. 71) records inflata Lam.
1811 (= bulbosa Bolten 1798) from Delagoa Bay; and brasiliensis Chemn. (!)
from Mozambique. Macnae & Kalk (1958. Nat. Hist. Inhaca Is., p. 128) add
elegans Lam. and scitula Marrat from Delagoa Bay.
Gen. SYLVANOCOCHLIS Melvill
1889. (Melvill) in Sowerby, 7. Conch., vi, p. 149 (subgen. Mariona, non Vayssiére 1879).
1903. Melvill. ibid., x, p. 325.
Shell ovate-fusiform, with thick walls. Columella concave, with parietal
callus constricting the posterior end of aperture, separated from outer lip by a
narrow slit, which is accentuated by the slight depression on last whorl forming
an indent in the outer lip. Canal short, wide. Last whorl with a spiral groove
near base forming a denticle on outer lip.
Operculum horny, large, oval, nucleus apical.
Radula with not very broad, tricuspid central plate, and bicuspid lateral
plate (as in Pseudoliva).
A monotypic, indigenous genus, if maintained distinct from Pseudoliva.
Sylvanocochlis ancilla (Hanley)
Fig. 15(a)
1859. Hanley. Proc. Zool. Soc. Lond., p. 429 (Pseudoliva a.).
1889. Sowerby. loc. cit., p. 149, pl. 3, fig. 2 (Pseudoliva a.).
1892. id. Mar. Sh. S. Afr., p. 15, pl. 1, fig. 14 (Pseudoliva a.) (this figure, reproduced from
the 1889 figure, does not show the characteristic posterior slit in the aperture).
1903 (1 July). Melvill. loc. cit., p. 324, fig. |
1903 (8 July). Sowerby. Mar. Invest. S. Afr., ii, p. 228 (Pseudoliva a.).
1904. Smith. 7. Malac., xi, p. 23 (Sylvanocochlea, err.).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 35 (Sylvanocochlea, err.).
1931. Tomlin. Ann. Natal Mus., vi, p. 430.
1932. ‘Turton. Mar. Sh. Port Alfred, p. 33 (? and pl. 7, no. 246).
ee
.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 61
Aperture 14-13 times the spire. Protoconch 13 whorls, alt. 1-5, diam.
2 mm., smooth (but worn). Postnatal whorls 4. Periostracum thick, forming
close axial wrinkles, the free margins of which are minutely fimbriate.
Operculum slightly convex, 23 * 12 mm. in 51 mm. shell.
Fulvous or slightly pinkish or violaceous, periostracum dull brown,
operculum dark brown. Animal pale, sprinkled with black specks around
margin of foot, siphon closely ringed with black.
Radula (of Algoa Bay specimen) with 57 rows, fig. 15(a).
aw W Tare eon
Re rw oe Ww
Fic. 15.
Central and lateral radula plates of (a) Sylvanocochlis ancilla (Hanley) ; (6) Ancilla bullioides Rve.;
(c) Melapium lineatum (Lam.). Central plate of (d) Ancilla_fasciata Rve.; (e) A. fasciata-albozonata
Smith; (/), (g) fasciata, variation of cusps; (hk) A. obtusa (Swainson) : (t) A. contusa Rve.;
(j) A. errorum Tomlin.
Kaffraria (Hanley), Port Elizabeth (Sowerby), Port Alfred (Bartsch,
Turton) ; off Great Fish River, 40 fathoms, one dead (S. Afr. Mus. P.F. coll.).
Living: 33°45 S. 26°44’ E. (off the Kowie), 40 fathoms (Sowerby 1903,
typ. err. 34° 45’ S. 25° 44’ E.); Algoa Bay, 39 metres, living (U.C.T.). Off
Cape St. Francis, 45 fathoms (Tomlin, fide Burnup).
The shell of the living specimen recorded by Sowerby from off the Kowie
is in S. Afr. Mus.; no report was published on the animal.
Gen. ANcCILLA Lam.
1799. Lamarck. Mem. Soc. H. N. Paris, i, p. 70.
1903. Won Martens. D. Tiefsee Exp., vii, pp. 37, 38.
1925. Thiele. ibid., xvii, p. 189 (S. Afr. Ancilla species, comments on von Martens 19093).
Shell ovate-fusiform, spire low or high, often covered with callus; body
whorl with basal spiral groove, which may form a denticle on outer lip. Parietal
callus often extending down the columella; canal short and broad. Operculum
oval, nucleus apical (or subapical); sometimes reduced in size or even absent.
Radula with tricuspid central plate, unicuspid lateral plate.
Remarks. A. osculata Sow. is transferred to the genus Bullia.
Although the basal groove is faintly traceable even in very worn specimens
of Ancilla, and would scarcely have been overlooked by Smith, one cannot help
62 ANNALS OF THE SOUTH AFRICAN MUSEUM
wishing for a re-examination of Bullia ancillaeformis Smith (1906. Ann. Natal
Mus., i, p. 37, pl. 7, fig. 8); the shell is so very like A. bullioides in shape. But
the locality (Natal) is against it being bulliozdes.
Ancilla obesa Sow.
Fig. 16( f )
1859. Sowerby. Thes. Conch., ili, p. 65, pl. 213, figs. 44, 45.
1932. Turton. Mar. Sh. Port Alfred, p. 32.
Not obesa von Martens 1903 = callifera Thiele 1925 = reever Smith.
Spire without any covering of callus, parietal callus thin, profile of sides
straight or very slightly concave, apex pointed. Aperture about twice as long
as spire. Postnatal whorls 4. Basal groove meeting columella slightly above
middle. 18 X 95 mm. Turton: 21 mm. long.
Operculum and radula?
Fawn, a spiral band of alternating orange-brown and white spots on upper
part of whorls, below this on body whorl an irregular series of spots, and
another series below the basal groove; between these, two series orange-brown
axial zigzags forming a reticulate pattern, lower part of base brown; reflected
columella white.
Port Elizabeth, Kowie and Port Alfred (Sowerby, Bartsch, Turton, S. Afr.
Mus.). Tongaat (north of Durban), East London, and Still Bay (S. Afr. Mus.).
The record of 6 specimens in S. Afr. Mus., registered as having come
from Hout Bay on west coast of the Cape Peninsula, is not acceptable.
No living specimens seen.
Ancilla reevei Smith
Fig. 16(e)
1903. Von Martens. D. Tiefsee Exp., vii, p. 37 (Ancillaria obesa, non Sow.).
1904. Smith. 7. Malac., xi, p. 29, pl. 2, fig. 11.
1925. Thiele. D. Tiefsee Exp., xvii, p. 190, pl. 33 (21), fig. 15, and text-fig. 7 (radula)
(callifera).
1932. Turton. Mar. Sh. Pt. Alfred, p. 31, and var. bipartita, p. 31, pl. 6, no. 233.
Spire covered with thick callus, especially above aperture, right profile
convex, left profile convex above, indented below (in the suture), apex pointed.
Aperture about twice as long as spire. Postnatal whorls 4 (Smith: ? 5). Basal
groove meeting columella about in middle. 22 x 10-11 mm. Turton: 24 mm.
long.
Operculum?
Fawn, spire mostly brown, a brown band between the sutural indent and
lower margin of callus with white spots on its lower border, a series of brown
spots above the basal groove, extending upwards as zigzag or undulate brown
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 63
axial lines, often joining to form a reticulate pattern; lower part of base with
2 brown bands and a white one between them; parietal callus and reflected
portion of columella white.
Radula (Thiele), central plate with small middle cusp flanked on either
side by a large cusp, and externally 4-5 denticles.
Port Alfred and the Kowie (Smith, Bartsch, Turton, S. Afr. Mus.).
34° 51'S. 19° 37’ E., 80 metres (von Martens, Thiele).
Fic. 16.
Ancilla, to show callus (stippled) and basal groove (a) bullioides Rve.; (6) contusa Rve.; (c) hasta
von Mariens; (d) errorum Tomlin; (e) reevet Smith; (f) obesa Sow.; (g) marmoratagRve.; (h)
Jfasciata Rve.; (i) obtusa (Swainson); (j) optima Sow. In the front view of obtusa the callus is
brown on the left, white on the right of the dotted line.
64 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. Thiele was correct in not accepting von Martens’s identification
of the Valdivia shell as obesa Sow., but he overlooked Smith’s species and
proposed a new name.
The indent on the left side of the spire is characteristic, and together with
the spire-callus distinguishes this species from obesa; the two species are very
similarly coloured and pattern alone is not always decisive in separating them.
The spire-callus is not an adult character, but occurs on younger examples
smaller than the largest ones of obesa; can it possibly be a sexual character ?
Turton’s bipartita lacks the indent on left profile of spire, but the specimen
appears to be water-worn.
Ancilla obtusa (Swainson)
Figs. 15(h), 16(2)
1825. Swainson. Q. 7. Scz., xviil, p. 282 (Ancillaria o.).
1848. Adams & Reeve. Voy. Samarang. Moll., p. 31, pl. 13, fig. 6.
1859. Sowerby. Thes. Conch., iii, p. 62, pl. 211, figs. 15, 16.
1886. Watson. Challenger Rep., xv, p. 229.
1913. Bullen Newton. Rec. Albany Mus., ii, p. 348, pl. 24, figs. 3, 4 (Baryspira sp.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 31.
Spire completely enveloped in callus. Shell ovate with shortly pointed
apex, or obovate with bluntly rounded apex. Aperture about 14 times spire.
Postnatal whorls ? (concealed in callus). Basal groove meeting columella at
posterior third. Callus spreading laterally to left at top of aperture in a broad
rounded lobe which is continued on to spire, and which is visible when shell
is viewed from back (more so in adult than in juv.). 47 X 28 mm. Turton:
54. mm. long.
Operculum oval, slightly concave, nucleus subapical, 15 x 8 mm. in
39 mm. shell with aperture 29 mm.
Fawn or buff, a warm brown band on the base; the same colour shows
through the callus on spire; callus on body whorl brown, parietal callus above
aperture white; often arcuate or flame-like streaks on the brown body-whorl
band; middle band margined above and below with white and usually showing
the growth-lines. Operculum yellowish-horny or amber. Animal (Mossel Bay,
U.C.T.) dirty white with rather coarse sepia-brown speckling.
Radula with 95-100 rows: middle cusp of central plate rather broad,
smaller than the other two, outer margins of the latter crenulate.
Fossil, Mio-Pliocene: Redhouse, near Port Elizabeth (Newton);
Quaternary: Sedgefield near Knysna (A. R. H. Martin).*
Port Elizabeth and Algoa Bay, Port Alfred (Sowerby, Bartsch, Turton,
S. Afr. Mus.). Tongaat (north of Durban), Still Bay, Hermanus (S. Afr. Mus.).
Simon’s Bay, False Bay, 15-20 fathoms (presumably living) (Watson). Mossel
Bay (living) (U.C.T.).
* For description of these deposits see: Martin. S. Afr. J. Sci., 52, p. 187, 1956.
4
CONTRIBUTIONS TO KNOWLEDGE OF 8.A. MARINE MOLLUSCA 65
Off Cape Natal (Durban), 62 fathoms, off Glendower Beacon (Port
Alfred area) and Great Fish Point, 66-100 fathoms, Mossel Bay and off Cape
St. Blaize, 17-19 fathoms, all dead (S. Afr. Mus. P.F. coll.).
Algoa Bay, and off East London, 10-20 fathoms, living (S. Afr. Mus.
Ea coll.).
Remarks. ‘Two young specimens 14 X 7 and 17 X 10 mm. are ovate in
shape with pointed apex, with slightly convex profiles subtending an angle a
little less than go°. Larger specimens (21 mm. upwards) may retain the pointed
apex (largest 42 mm.), or show various stages in its envelopment in callus, the
final stage being a blunt and broadly rounded apex.
Four juveniles 2-5-4 mm., and one 6 mm. long (Still Bay, Muir coll.) may
be this species, but no connecting links between the 6 and 14 mm. specimens
are available. The smallest is 2-5 x 1:3 mm. 14 whorls. The 6 mm. specimen,
though slightly worn, shows the characteristic shape of the callus. Juveniles
7 mm. and upwards of fasciata-albozonata (a species common at Still Bay) have
a narrower apex, and the callus does not expand laterally in a rounded lobe.
Ancilla optima Sow.
Fig. 16(j)
1892. Sowerby. Mar. Sh. S. Afr., p. 16 (australis, non Sow.).
1897. id., Append. Mar. Sh. S. Afr., p. 7, pl. 6, fig. 31.
Shell ovate, spire not completely enveloped in callus, apex pointed, both
profiles nearly straight. Aperture 14-12 times spire. Postnatal whorls ? Body
whorl gently shouldered, on upper portion with growth-lines forming low
arcuate, obliquely protractive lirae. Basal groove meeting columella at about
posterior quarter. Parietal callus expanding laterally, but not in a broad
rounded. lobe, and not visible when viewed from back, scarcely reaching apex.
Boe 2 7 MOM.
Operculum oval, slightly concave, nucleus subapical, 17 X 9 mm. in
37 mm. shell with aperture 21 mm.
Fawn or buff; upper part of body whorl, spire, and base chestnut-brown,
the upper brown band sharply defined from the pale middle band (which
includes the basal groove) but becoming paler towards the (concealed) suture,
the arcuate lirae paler than the ground colour.
Radula with about 75 rows (juv.); middle cusp of central plate narrow,
shorter than the other two, outer margin of the latter denticulate, a minute
denticle on either side at base of middle cusp (cf. fig. 15(b), bulliozdes).
Durban (Sowerby); Pondoland coast (S. Afr. Mus.); off Durban,
40 fathoms, from fish stomachs (S. Afr. Mus. Ross-Frames coll.).
Living: off Cape Vidal and O’Neil Peak (Zululand), off Cape Natal
(Durban), and off Cape Morgan, 40-80 fathoms (S. Afr. Mus. P.F. coll.).
Delagoa Bay (U.W.).
66 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. The arcuate lirate growth-lines are distinctive.
The Delagoa Bay specimen is very dark, especially the base and the callus.
The lirate growth-lines are covered by a film of callus and indistinct.
Three specimens from Tongaat (30 miles north of Durban) are also
rather dark, the middle band darker than in typical optzma.
Ancilla contusa Rve.
Figs. 15(z), 16()
1864. Reeve. Conch. Icon., pl. 9, fig. 31.
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 7, pl. 6, fig. 23 (decipiens).
1903. id. Mar. Invest. S. Afr., ii, p. 228, pl. 3, fig. 3.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 34 (decipiens).
Shell ovate, apex pointed, spire completely (or nearly so) enveloped in
callus, profiles nearly straight or slightly convex, subtending an angle of
c. 55°-60°. Aperture a little longer than (up to 1} times) spire. Postnatal
whorls ? Basal groove meeting columella nearly at top of aperture. Callus
expanding slightly at suture of body whorl (cf. optima) and continued on to
spire. 52 X 25 mm.
Operculum oval, slightly concave, nucleus subapical, 11 X 5:5 mm. in
25 mm. shell with aperture 14 mm.
Cream or pale buff, spire usually white, but a faint brown sutural band
sometimes shows through the callus; upper brown band usually well marked,
especially its lower edge (where callus ends); a chestnut-brown band on base;
on middle of body whorl a series of spiral brown bands or lines, varying in
number and intensity, sometimes very indistinct.
Radula with ¢c. 95 rows, middle cusp of central plate rather broad but
smaller than the other two, outer margin of latter crenulate or denticulate,
one or two minute denticles on either side of base of middle cusp.
Durban and Natal, 25-30 fathoms (Sowerby); Port Elizabeth (Sowerby:
decipiens); Port Alfred (Bartsch: decipiens); Pondoland coast (S. Afr. Mus.).
Natal, 25-27 fathoms, and from fish stomachs off Durban (S. Afr. Mus.
P.F. coll.). Durnford Point (Zululand) 13 fathoms, living (S. Afr. Mus.
P.F. coll).
Remarks. A. decipiens is the beach-worn form of contusa tinged with pink
or violaceous, as are several other South African beach-weathered shells.
Some specimens, especially when the spiral brown bands are indistinct
or bleached out, may be difficult to separate from optima, but the arcuate lirae
of the latter are usually distinctive.
Ancilla bullioides Rve.
Figs. 15(6), 16(a)
1864. Reeve. Conch. Icon., sp. 37.
1886. Watson. Challenger Rep., xv, p. 229 (montrouzieri, non Souv.).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 67
1903. Sowerby. Mar. Invest. S. Afr., uu, p. 228 (bulloides [sic]).
? 1903. Von Martens. D. Tiefsee Exp., vil, p. 38 (dimidiata, non Sow. part: Sta. 1193).
?1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 35 (bulloides [sic]).
1925. Thiele. D. Tiefsee Exp., xvii, p. 191, pl. 33 (21), fig. 19 (cf. montrouzieri).
? 1932. Turton. Mar. Sh. Pt. Alfred, p. 32 (bulloides [sic]).
Bullia-like, the greatest width anterior to (below) middle of length.
Ovate, spire high. Aperture a little longer than spire, expanded in lower half.
Postnatal whorls ? Basal groove meeting columella nearly at top of aperture.
Callus expanding laterally over the suture, extending to and covering the apex
(the actual! point of the apex is only thinly covered), partly visible in back view.
40 X 17 mm.; smallest example seen 14°5 mm. long.
Operculum slightly concave, oval, narrowing above, inner margin
sinuous, nucleus subapical, 17 <X 7 mm. in 40 mm. shell with aperture 20 mm.
Creamy-white, spire pale pinkish-orange or buff, a spiral band of the same
colour on base. Operculum yellowish-horny or amber.
Radula with c. 75 (juv.) to 95 (adult) rows; middle cusp of central plate
narrow, smaller than the other two, sometimes markedly so, outer margin of
latter crenulate or feebly denticulate.
Off Cape of Good Hope, 98 fathoms (Watson); off Cape Peninsula,
190 fathoms (Sowerby). Living: south of False Bay, around Cape Point, off
west coast of Cape Peninsula, and northwards to the Saldanha Bay area,
85-195 fathoms (S. Afr. Mus. P.F. coll.).
35° 11'S. 23° 2’ E., 500 metres, dead shells inhabited by Hermit crabs
(Thiele: Sta. 103, fig. 19).
The identification of specimens from Port Alfred (Bartsch, Turton) is very
doubtful.
Remarks. Watson had one specimen and was unable to find any out-
standing differences between it and the New Caledonian montrouzieri. Thiele
also compared some specimens with the latter species. Nevertheless the two
species are not likely to be synonymous. It is even more unlikely that this deep-
water Cape species should be ‘common’ (Turton) on the beach at Port
Alfred!
I have seen no authentic dimidiata. Von Martens recorded this species
from three localities. I am inclined to think that his specimens from Sta. 113,
south of False Bay, 318 metres, were really bullioides; those from shallow water
70 metres in Simon’s Bay (False Bay) Sta. 114 were errorum; the third lot from
Sta. 104 are from a locality farther east, not far from where hasta has been
obtained.
Thiele suggested identifying all these with bullioides; but his fig. 18 is
certainly not bulliozdes.
On the other hand, the specimens covered with Epizoanthus and inhabited
by the Hermit crab Parapagurus pilosimanus, which Thiele compared with
montrouziert, are undoubtedly bulloides, although his fig. 19 has not quite the
characteristic shape.
68 ANNALS OF THE SOUTH AFRICAN MUSEUM
In the Preter Faure collection there are
numbers of such specimens. The Hermit
crab has a wider distribution around the
southern slope of the Agulhas Bank, and
as far east as off East London (Barnard.
Ann. S. Afr. Mus., xxxvill, p. 451, 1950),
but utilizes any dead shell. Its distribution
is therefore no criterion for the distribution
of the mollusc; in fact the only specimens
of the crab inhabiting Ancilla shells come
from the Cape Point area where living
bullioides have been obtained. Thiele’s
record was 35° 11’ S. 23° 2’ E., 500 metres.
The Epizoanthus colony on a dead
shell inhabited by Parapagurus continues
to grow in a helicoid spiral, the pitch
increasing pari passu with the girth of the
crab. When a crab has occupied and out-
grown an Ancilla shell the helicoid
te a2 addition does not continue at the sharply
Section of Ancilla shell and the Coelente- acute angle of the Ancilla shell, but at a
rate Epizoanthus inhabited by Hermit- wider angle, so that in external view the
crab (Parapagurus pilosimanus), showing ; Sas
alteration in angle of spiral when the crab appears to be occupying a Buccinoid
crab has outgrown the Ancilla shell. shell (tie.) 17):
Ancilla errorum Tomlin
Figs. 15(j), 16(d)
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 229 (angustata, non Sow.).
? 1903. Von Martens. D. Tiefsee Exp., vii, p. 38 (dimidiata, non Sow., part: Sta. 114).
1906. Smith. Ann. Natal Mus., i, p. 28 (Marginella angustata).
rg21. “Tomlin. 7. Conch... xvi, p: 216; plo, mph.
Shell ovate, greatest width at middle length. Spire moderately high,
enveloped in callus. Aperture a little longer than spire. Postnatal whorls ?
Basal groove meeting columella at upper quarter of aperture. Callus expanding
laterally and continued almost to apex, not visible in back view. 15 * 7 mm.
Operculum oval, slightly concave, nucleus subapical, 6:5 x 2 mm. in
shell 14 mm. with aperture 8 mm.
Pale buff or fawn, a brown sutural band, and another band on base.
Operculum yellowish-amber.
Radula with c. 80 rows, middle cusp of central plate smaller than the
side cusps, outer margin of latter denticulate, a minute denticle on either side
at base of middle cusp.
Off Cape Point, 42 fathoms (Sowerby, Tomlin).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 69
False Bay, Walker Bay, Algoa Bay, and off Umhloti River (Natal),
20-47 fathoms (S. Afr. Mus. P.F. coll.). 35° S. 20° 49’ E., 91 metres (s.s.
Africana, per U.C.T.).
Remarks. Smith stated that Sowerby inadvertently recorded this species as
an Ancilla instead of a Marginella. The statement, however, is incorrect and
should be ignored, because the specimens recorded by Sowerby and returned
by him are in fact Ancilla. The same specimens were later seen by Tomlin, who
also saw the Natal specimen recorded above.
On account of the locality I am inclined to think the Valdivia specimens
from Sta. 114, Simon’s Bay (False Bay), 70 metres, identified by von Martens
as dimidiata, should be referred here. Thiele without any mention of 'Tomlin’s
species, referred them to bulliozdes.
The species is a small edition of bullioides living in shallower water. A
juvenile bulliordes, 14:5 mm. long, however, is easily distinguished by the shape
of its shell and of the callus.
Ancilla hasta von Martens
Fig. 16(c)
1902. Von Martens. SB. Ges. naturf. Fr. Berlin, p. 241.
Eeoeeretees DD). Wrefsee Hxp., vil, p. 37, pl. 3, fig. 13.
? 1903. id. ibid., p. 38 (dimidiata, non Sow., part: Sta. 104).
?1925. Thiele. ibid., xvii, p. 191, pl. 33 (21), fig. 18 (bulloides, non Rve.).
Shell fusiform, greatest width at middle length, spire high, apex pointed.
Aperture about 14 times spire. Postnatal whorls 7 (von Martens). Basal groove
meeting columella at upper third of aperture. Callus expanding on body
whorl suture, extending to but not covering the preceding suture, not visible
in back view. 25 X 10 mm. Three other specimens 16-19 mm. long.
30 X 11 mm. (von Martens).
‘Pale rosy-fulvous, white towards apex’ (von Martens). Creamy-white,
callus and sutures slightly darker buff.
35° 10'S. 23° 2’ E., 500 metres (von Martens).
65-73 miles off Cape St. Blaize, 85-90, and 125 fathoms (S. Afr. Mus.
Ese eoll.).
Remarks. Von Martens does not mention the shape and extent of the
callus. The Pieter Faure localities are near to that of the Valdivia, and the four
dead specimens taken by the former vessel are undoubtedly referable to hasta.
The Valdivia specimens from Sta. 104. 35° 16'S. 22° 26’ E., 155 metres,
recorded by von Martens as dimidiata, may possibly be hasta. Thiele’s figure 18
is much more like a hasta than a bullioides.
Ancilla fasciata Rve.
Figs. 15 (d-g), 16(h)
1864. Reeve. Conch. Icon., sp. 44.
1892. Sowerby. Mar. Sh. S. Afr., p. 16 (cinnamomea, non Lam.).
70 ANNALS OF THE SOUTH AFRICAN MUSEUM
1904. Smith. 7. Malac., xi, p. 29, pl. 2, fig. 9 (albozonata).
1925. Thiele. D. Tiefsee Exp., xvii, pp. 190, 191, pl. 33 (21), fig. 17 (agulhasensis).
1932. Turton. Mar. Sh. Pt. Alfred, p. 32, pl. 6, no. 238 (pattern var.) and p. 32 (albozonata).
f
}
‘
Aperture about twice the spire. Basal groove meeting columella near
upper end of aperture. Callus beginning about midway on columella, not much
expanded laterally, continued upwards nearly to suture with preceding whorl.
The lower of the two incised lines on base (the upper being the basal groove)
nearer to the latter than to the reflected columella (contrast marmorata).
26 h12 am,
Operculum present but reduced in size, ovoid-oblong, slightly concave,
growth-lines distinct, with very fine striae radiating from nucleus, 3 X 2 mm.
in 17 mm. shell with 11 mm. aperture.
Brown, apex white, a white band below the brown suture on body whorl,
another white band above reflected columella on base. The variety figured by
Turton chequered with oblong brown spots.
Animal pale biscuit colour.
Radula with 80-90 rows, cusps on the central plate variable: all three may
be subequal in size, with 1-4 denticles on either side of the middle cusp; some-
times there are (3) 4-5 cusps between the large side cusps, varying in size and
arrangement, sometimes two of them united to form a bifid cusp. Thiele found
the central plate had 5 smaller, somewhat irregular cusps between the two big
ones.
Port Elizabeth and Port Alfred (auct. et S. Afr. Mus.).
St. Francis Bay, 80-100 metres, Algoa Bay. 35° 16'S. 22° 26’ E.,
155 metres, and (living) Simon’s Bay (False Bay), 70 metres (von Martens,
Thiele).
Off Tugela River (Natal), 37 fathoms, 1 alive; off Umhloti River (Natal),
100 fathoms, 1 dead; off Cape Natal (Durban), 85 fathoms, 1 alive; Algoa Bay
to off Walker Point, 37-66 fathoms, living (S. Afr. Mus. P.F. coll.). False Bay,
living (U.C.T.).
Remarks. As typical of this species and of marmorata I have had for com-
parison examples identified by Tomlin.
Thiele was unable to find any trace of an operculum in his specimens.
His description of the radula bears out the variability of the cusps which I have
found in several examples.
Ancilla marmorata Rve.
Fig. 16(g)
1864. Reeve. Conch. Icon., sp. 32.
1892. Sowerby. Mar. Sh. S. Afr., p. 17, pl. 1, fig. 15 (pura).
1903. Von Martens. D. Tiefsee Exp., vii, p. 38.
1906. Smith. Ann. Natal Mus., i, p. 27, pl. 7, fig. 4. (ordinaria).
1925. Thiele. D. Tiefsee Exp., xvii, p. 190, pl. 33 (21), fig. 16 (referred with ? to agulhasensis).
1932. Turton. Mar. Sh. Pt. Alfred, p. 32, and var. major.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 71
Distinguished from fasciata by the two incised lines on base being farther
apart, the lower one being midway between the upper line (which is the basal
groove) and the reflected columella; clearly shown in Smith’s and Thiele’s
figures. 19 X 9 mm. (S. Afr. Mus.), 21 X 9 mm. (Sowerby); 20 mm. long
(Turton: var. major).
Operculum present but reduced in size, ovoid, with distinct growth-lines
and very fine radiating striae, 2-5 X 1:5 mm. in Io mm. shell with 6-5 mm.
aperture.
Fawn or pale buff, sometimes with faint mottling or axial lines, a darker
sutural band on body whorl, and another between the two incised lines; or
pure white.
Radula with about 70 rows, cusps on central plate subequal in size, no
denticle on either side of middle cusp (only one specimen examined).
Port Elizabeth (Sowerby: pura); Port Alfred (Turton, S. Afr. Mus.) ;
Port Shepstone (Natal) (Smith: ordinaria).
Bato o 22 26 H.. 155 metres (von Martens, Thiele).
Off Umhloti River (Natal) 40 fathoms, dead; off Cape Natal, 54 fathoms,
living; off Umkomaas (Natal), 40 fathoms, living; off Cove Rock (East London
area), 22 fathoms, dead but fresh; off Glendower Beacon (Port Alfred area),
66 fathoms, 1 alive; False Bay, 32 ‘thome Io alive (oy str Vlas? EE coll:
False Bay, 1 alive (U.C.T.).
Remarks. A. pura appears to be a somewhat slender example of this species,
but Sowerby’s figure is inadequate.
One living specimen was taken by the Preter Faure off the west coast of the
Cape Peninsula in 60 fathoms; it is white, and seems indistinguishable from
marmorata. 1 should have assumed an error in labelling, unless confirmation
had been provided by finding living examples in an intermediate locality, viz.
False Bay (P.F., and U.C.T.). Its radula has 60 rows, the 3 main cusps on
central plate subequal, on one side of the median cusp one small cusp, on the
other side 2 small cusps.
The P.F. specimen from False Bay is fawn coloured, but the U. C. Es
example is white.
Gen. Metapium H. & A. Adams
1853. H. & A. Adams. Gen. Moll., i, p. 136.
1889. Smith. Ann. Mag. Nat. Hist. (6), ii, p. 267.
1929. Thiele. Handbuch, i, p. 332.
1937. Peile. Proc. Mal. Soc., xxii, p. 182 (radula).
Shell broad, rounded-piriform, body whorl bulbous, outer lip expanded
in adult. Spire low, protoconch papilliform. Columella curved, with a ridge
from middle bordering the canal; strong parietal callus within aperture
posteriorly. Periostracum thin. No operculum.
Radula with broad central plate, tricuspid, lateral plate unicuspid (as in
Oliva).
72 ANNALS OF THE SOUTH AFRICAN MUSEUM
Melapium elatum (S. & W.)
1829. Schubert & Wagner. Conch. Cab., xii, pp. 92, 94, pl. 226, figs 4012, 4013 (Pyrula e.).
1go1. Smith. 7. Conch., x, p. 110.
1952. Bayer. ool. Med., xxxi, no. 25, p. 297 (as syn. of lineatum).
Large. Whorls 4, last whorl with a blunt but definite shoulder.
63 X 61 mm. (protoconch worn).
White with numerous narrow axial brown or orange lines, slightly
undulate and sometimes broken into discontinuous streaks.
Off Durban, 40 fathoms (from fish stomachs) (Smith, 1gor).
Distribution. East Indian seas.
Remarks. Smith’s record is the only one with precise locality. S. Afr. Mus.
has 4 large specimens and one half-grown from the Ross-Frames collection,
without locality; but they look as if they might have been taken from fish
stomachs, from which source Ross-Frames is known to have obtained some of
his shells.
Bayer includes this species as a synonym of lineatum. Probably only one
species should be recognized. The smallest of the 5 specimens ex coll. Ross-
Frames measures 40 < 38 mm. and has a definite shoulder on the last whorl.
Melapium lineatum (Lam.)
Fig.15(c)
1822. Lamarck. Anim. sans. Vert., vii, p. 147, no. 27 (Pyrula 1.).
1856. Wood. Index Test., ed. Hanley, p. 212, pl. 4, fig. 8 (Buccinum bulbus).
1889. Smith. Ann. Mag. Nat. Hist., ui, p. 269 (with radula).
1903. Von Martens. D. Tiefsee Exp., vii, p. 26.
1903. Thiele. ibid., p. 166, pl. 9 (4), fig. 51 (radula).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 62 (bulbosum err.).
1925. Thiele. D. Tiefsee Exp., xvii, p. 191.
1932. Turton. Mar. Sh. Port Alfred, p. 79 (bulbosum err.).
1937. Peile. loc. cit., p. 182, fig. 16 \(radula):.
1951. Barnard. Beginner’s Guide S. Afr. Sh., p. 69, pl. 6, fig. 15 and fig. 37 (egg-capsules).
1952. Bayer. <ool. Med., xxxi, p. 297 (references).
Protoconch, all specimens worn. Postnatal whorls 4. Last whorl without
any trace of a shoulder; 3rd and beginning of 4th whorl with 4-5 feeble spiral
striae on upper part. 30 X 28 mm.; smallest example seen 9 X 6 mm.
White with numerous axial brown or orange lines, sometimes undulate or
arcuate or discontinuous, sometimes in addition an irregular series of oblong
(axially) spots around the periphery; anterior canal externally suffused with
crimson.
Animal (as preserved) foot dull buff with 2 submarginal red lines;
Prof. Day says the margin is blue when alive. —
Radula with 60-65 rows.
Natal (Krauss); Port Alfred, Port Elizabeth and Algoa Bay, St. Francis
Bay (auct.); Port St. Johns and Still Bay (S. Afr. Mus.); O’Neill Peak (Zulu-
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 73
land), 90 fathoms; off Tongaat, 36 fathoms (S. Afr. Mus. P.F. coll.). Living:
East London to Mossel Bay, 19-50 fathoms (S. Afr. Mus. P.F. coll.), also
Mase! Bay (U.C.T.); 33° 5. 28° 11’ E., 31 fathoms (U.C.T.).
Remarks. Some of the Still Bay beach specimens (Muir coll.) are quite
fresh, with unfaded brown streaks and spots; quite possibly the species lives as
far west as this locality.
Egg-capsules. Six specimens (P.F. coll. Nov.-Dec. 1898, off Gape Recife)
each with 2 capsules attached on outer side of the parietal wall and curving
inwards towards the aperture. 10-11 X 5-6 mm., subcylindrical, cross-section
somewhat triquetral; the two inner surfaces horny, but the outer surface feebly
calcified. One contained a single crushed embryo; all the others were longi-
tudinally split and empty. This embryo could not be definitely determined as
a Melapium, consequently these capsules can only be presumed to belong to this
mollusc. It seems unlikely, however, that another species would deposit its
egg-capsules in this position, and moreover they have not been found attached
to any other Gastropod.
One of the U.C.T. East London specimens also carried two egg-capsules
(empty): 21 May 1958.
Fam. FASCIOLARIDAE
Gen. FAscIOLARIA Lam.
1g1t. Strebel. 7B. Hamb. Wiss. Anst., xxviii, pp. 1-58, 15 pls.
Shell usually large, fusiform, not umbilicate; canal rather long. Parietal
callus present or absent; columella with 3 pleats, sometimes with additional
wart-like nodules; columella glaze adnate throughout to the rostrum. Proto-
conch large.
Operculum ovate, apex incurved, nucleus apical; internal surface with
thickened shiny margin, especially at apex.
Radula long, with 3 plates in each row, central plate quadrangular,
tricuspid (sometimes a smaller one on either side externally), lateral plate wide,
with numerous subequal cusps; the number of the latter increases with growth
and is not always a specific character.
Ege-capsule stalked.
Remarks. As in many other genera plump and slender individuals of the
same species occur. Strebel (pp. 1, 42) mentioned the possibility of this being
a sexual difference, and deplored the lack of evidence. Although only 5
individuals of one species of this genus have been available to me for sexing,
they gave no support to the suggestion. (cf. Burnupena delalandii, p. 166).
Key to the South African species
PeRpARiCrM eae et 5. es Sk see eee 8? amentose
2. Parietal callus present.
74 ANNALS OF THE SOUTH AFRICAN MUSEUM
a. Profile convex, without angular shoulder.
i, Spiral sculpture|fime and regular . > 42 4 : “| a yh coneneieag
ii. Spiral sculpture irregular, coarser and finer lirae more or es alter-
Mating. 2 eee, ee ee ee og awe
b. Profile angular, with shoulder knobs.
i. Protoconch small, diam. 1:5 mm. Shell usually with dark ee
lines, usually in pairs, sometimes unicolorous. . . . . trapezium
ii. Protoconch large, diam. (2°5 mm. worn) 3°5-5 mm. Shell uni-
colorous 2 ee ee ee ee
Fasciolaria filamentosa Lam.
1880. Von Martens. Mauritius G Seychellen, p. 245.
1911. Strebel. loc. cit., p. 34, pl. 6, figs. 33, 34; pl. 7, figs. 35-37; pl 15, ieee
1929. Thiele. Handbuch, i, fig. 377 (radula).
1952. Satyamurti. Bull. Madras Govt. Mus., n.s. 1, 2, pt. 6, p. 185, pl. 17, figs. 9 a-c.
1952. Braga. Anais Est. Zool. Invest. Ultramar., vii, 3, p. 73, pl. 2, fig. 2.
No parietal callus. Aperture longer than spire. Profile convex, sometimes
with slight shoulder. Protoconch 14 whorls, small (see Strebel, pl. 7, fig. 37),
smooth. Postnatal whorls 7; 1st with 7 axial ribs, increasing to 11 on middle
whorls, on later whorls forming only slight swellings on the shoulder; crossed
by 4 spiral lirae on 1st whorl, increasing in number on middle and later whorls,
more or less regular above shoulder, but alternately broad and narrow below,
the larger ones becoming better marked on the base. Columella pleats distinct,
glaze narrow; outer lip internally plicate. 172 X 71 mm. (Strebel).
Radula, lateral plate with 13 cusps (Thiele, fig. 377).
Natal (Sowerby), Mozambique (Braga).
Distribution. Red Sea, Zanzibar, Réunion, Mauritius, Seychelles,
Madagascar and Indo-Pacific to Japan.
Remarks. A variable species (Strebel), but distinguished by the absence
of the parietal callus.
Turton’s record from Port Alfred (1932. Mar. Sh. Pt. Alfred, p. 49) is
probably not filamentosa. Sowerby’s Natal record was probably a dead shell.
Fasciolaria rutila Watson
Figs. 18(a), 19(c)
1882. Watson. 7. Linn. Soc. Lond., xvi, p. 335.
1886. id. Challenger Rep., xv, p. 242, pl. 13, fig. 6.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 227, pl. 3, fig. 2 (juv. shell and radula).
1903. Von Martens. D. Tiefsee Exp., vii, p. 30.
? 1923. Odhner. Géteb. K. Vet. Handl., xxvi, p. 6 (= Med. Géteb. Mus., xxiii).
? Tomlin. Ann. Natal Mus., v, p. 290.
Thin-walled, profile of whorls convex, without any shoulder. Protoconch
(uncorroded) large, subglobular, 1 or 14 whorls, diam. 5:5—7 mm., alt. 5-6 mm.
(corroded: diam. 3°5, alt. 3 mm.). Postnatal whorls 5, smooth in appearance,
but with numerous fine spiral lirae closely set, c. 11-12 on 1st whorl, becoming
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 75
a little stronger and more widely spaced on later whorls. Parietal callus
present; columella with 3 pleats, but posterior one feeble and obscure in large
specimens. Canal long, narrow, sigmoid; outer lip thin, not plicate internally.
Periostracum thin. 132 X 51 mm.
Operculum 35 X 19 mm. in 132 mm. shell.
White with pale yellowish brown periostracum, operculum dark brown.
Radula with 230-250 rows, usually an accessory denticle externally on
both sides on the central plate; lateral plate with 8 cusps in juv. (figured by
Sowerby), in adult 13 on one side and 14 on the other (2 specimens), or 15 and
16 resp. (one specimen) (excluding the tiny internal denticle).
Fre. 1G:
(a) Fasciolaria rutila Watson, protoconchs of two shells.
(6) F. lugubris Rve., two views of protoconch extracted from
(c) egg-capsule. (d) F. heynemanni Dnkr., protoconch.
imine: 95°45. 18° 37 E., 150 fathoms (Watson); 33° 41 S. 18° E.,
178 metres (von Martens); off Cape Peninsula, 154 fathoms (Sowerby: juv.);
off Cape Point, 85-145 fathoms (S. Afr. Mus. P.F. coll.); 33° 11'S. 17° 57’ E.,
77 metres; 31° 39’ 8. 16° 55’ E., 287 metres (s.s. Africana) ; ? off Umhloti River
(Natal), 40 fathoms (Sowerby).
Dead? St. Sebastian Bay, 40 fathoms (Odhner); off Cape St. Francis
(Tomlin, coll. Burnup).
Remarks. ‘The Burnup shell recorded by Tomlin was 5} in. (133 mm.) long.
The Pieter Faure obtained only one shell off the Natal coast (if the label was
correct); all the others were from off the Cape Point. The Africana has shown
that the distribution extends to 31° S. on the west coast. The large Natal
76 ANNALS OF THE SOUTH AFRICAN MUSEUM
specimen, now in the British Museum, should be re-examined on account of
its alleged locality.
More probable misidentifications are Odhner’s 150 mm. and Burnup’s
133 mm. shells. I consider that both of these are more likely to be the off-shore
form of lugubris, which has stronger and more irregular spiral liration than
rutila; the latter also has a more sigmoid canal.
Fasciolaria lugubris Rve.
Figs. 16(0, ¢), 19(@, 0)
1847. Reeve. Conch. Icon., sp. 2.
1848. Krauss. Stidafr. Moll., p. 110, pl. 6, fig. 12 (badia).
Toit. Strebel: loc: cit.,.p./ 9, pl) 6, tias.30,, 304, 31-
1932. Tomlin. Ann. S. Afr. Mus., xxx, p. 157, fig. 1 (agulhasensis).
1932. Haughton. Tr. Geol. Soc. S. Afr. (1931), pp. 34, 49-
Aperture longer than spire. Profile of whorls evenly convex, but sometimes
with faint indication of a blunt shoulder, rarely a definite shoulder with low
knobs. Protoconch (from egg-capsule) 14 whorls, diam. 3—3°5, alt. (i.e. apex
to rostrum) 5 mm., smooth, later part of whorl with 6-8 feeble axial ribs,
crossed by spiral lirae, c. 20 from suture to lower end of outer lip. Postnatal
whorls 7; 10 axial ribs on 1st whorl, on 2nd and 3rd and sometimes 4th a
similar number of slight undulations or low rounded knobs, obsolete on later
whorls; crossed by numerous spiral lirae, varying in size, usually coarser and
finer alternating, the larger ones sometimes with one or more striae, the peri-
pheral lira sometimes distinctly stronger than its neighbours, almost a costa
(agulhasensis), sometimes a pair of peripheral lirae stronger than the others.
Parietal callus present, but weak in young and half-grown shells; columella
pleats distinct. Outer lip evenly convex, not so distinctly incurved at beginning
of canal as in heynemanni, plicate internally, edge at some stages of growth
denticulate. Periostracum fibrous-fimbriate. 183 (tip of canal broken, about
5 mm. missing) * 75 mm. (type of agulhasensis); two False Bay examples (tip
of canal broken in both) 145 « 52 mm. and 142 x 55 mm. Largest littoral
example 81 (tip of canal broken) x 43 mm.
Operculum 28 X 14 mm. in 81 mm. shell.
Protoconch and first 2 (24) whorls white, rest fulvous or chestnut-brown,
operculum dark brown. Animal sealing-wax red with white dots on sides of
foot (ie:EL. Ba)
Radula: from protoconch in egg-capsule, 40 rows, lateral plate with 6
cusps; from 18 mm. shell 135 rows, lateral with 8 cusps; from 33 mm. shell
190 rows and 11—12 cusps resp.; from 85 mm. shell 225 rows and 12 cusps resp. ;
from 142 mm. shell 265 rows and 14-16 cusps resp. (in all cases excluding the
tiny internal denticle); central plate quadrangular, with well-developed cusps,
the middle one stronger than the side cusps.
Egeg-capsule club-shaped, 20 mm. high, distally elliptical in section, major
diam. 9 mm., minor diam. 8 mm., horny; only one specimen seen, containing
_ CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 77
4 embryos, but the apex was open and some of the embryos had probably
escaped.
Fossil, late Tertiary; Saldanha Bay (Haughton).
Dead. Natal (Krauss); False Bay (Strebel, coll. Fritsch); Cape Hangklip
(S. Afr. Mus.); between Cape St. Blaize and Flesh Point, 28 fathoms (S. Afr.
Mus. P.F. coll. type of agulhasensis).
Living. Steenberg Cove (St. Helena Bay) and Langebaan (Saldanha Bay),
littoral (U.C.T.); Oudekraal, west coast of Gape Peninsula, littoral (U.C.T.);
Sea Point, Cape Town, littoral (S. Afr. Mus.); Kalk Bay (False Bay), littoral
(S. Afr. Mus.), and 4-5 metres (U.C.T.).
Remarks. The striae on the larger spiral lirae are not (in the specimens I
have seen) so prominent as Strebel described in his specimens, and cannot be
regarded as a specific character.
I have seen a 51 mm. shell from Cape Hangklip (S. Afr. Mus. no. 14052),
and a 95 mm. shell from False Bay (U.C.T.) with distinct angular shoulders and
two peripheral lirae. Another False Bay example 142 mm. long (U.C.T.) has
g peripheral swellings on the 6th whorl, but none on the preceding whorls.
The resemblance of these specimens to heynemanni is strong, but the latter species
has finer and more regular lirae above the shoulder, and the shoulder pro-
jections are definitely knobs, not mere swellings or undulations; and the 2
peripheral lirae are situated in the middle of the whorl, not as in the dunkert
form of heynemanni adjoining the suture of the following whorl.
The contrast between the smooth slender examples and, e.g. the
95 X 46 mm. shouldered specimen is very striking, and is paralleled with the
adamsu and ocelliferus forms of Fusus verruculatus (p. 90).
Littoral examples are smaller and stouter than off-shore forms (cf.
heynemannt).
The 145 X 52 mm. shell from False Bay, 23 fathoms, was identified by
Tomlin as filamentosa in spite of its having a parietal callus. It bridges the gap
in size between the common littoral form and agulhasensis. The latter was
contrasted by Tomlin with scholvient, an obviously different species, but he did
not mention any differences between his n. sp. and lugubris. In fact there are
no differences.
Krauss said his badia was very like Fusus mandarinus, and also bore resem-
blance to Fasciolaria fusiformis and filamentosa.* ‘There are no later records of
lugubris from Natal, or indeed from east of Cape St. Blaize.
Fasciolaria trapezium Linn.
1880. Von Martens. Mauritius G Seychellen, p. 245.
1895. Cooke. Cambr. Nat. Hist., ii, fig. 121 (radula).
1903. Smith. Proc. Mal. Soc., v, p. 368 (heynemanni, non Dnkr.).
* See also Fusivoluta pyrrhostoma (Volutidae) for the Gazelle shells reported to be Fusus
mandarinus (p. 30).
78 ANNALS OF THE SOUTH AFRICAN MUSEUM
1911. Strebel. loc. cit., pp. 40 sqq. and vars. pls. 7-10, 13, 14, figs. 38-45, 48, 49, 61, 62.
1930. Fulton. Ann. Mag. Nat. Hist. (10), vi, p. 685, pl. 18, figs, 2, 2a (strebelz).
1952. Satyamurti. Bull. Madras Govt. Mus., I, 2, pt. 6, p. 186, pl. 17, fig. 10.
1952. Braga. Anais Est. Zool. Invest. Ultramar, vii, 3, p. 73, pl. 2, fig. 1.
Aperture longer than spire. Profile of whorls with angular shoulder.
Protoconch 14 whorls, diam. 1-5 mm., smooth (Strebel, figs. 38, 42). Postnatal
whorls 9 (? 10); spiral grooves only on early whorls but indicated on later
whorls by dark lines; on 1st and and (—3rd) whorls 8-9 axial ribs extending
from suture to suture, but on later whorls gradually restricted to the periphery
and forming knobs on the shoulder, which becomes definitely marked from 4th
whorl onwards: 5-7 in forma typica, 8-11 in varieties. Parietal callus present,
but weak in juveniles. Columella with 3 pleats, and often additional wart-like
nodules anterior to the main pleat; canal straight or slightly sigmoid; outer lip
internally with numerous plicae, usually a smooth zone between the plicae and
the edge; the latter in some stages of growth denticulate. Up to 215 mm. long
(f. typica); var. ponderosa 230 mm. (Strebel).
White or pinkish, with narrow brown spiral lines usually in pairs, plicae
in aperture orange-brown, periostracum yellowish-brown.
Radula (only a portion removed from a 153 mm. shell); lateral plate with
30 cusps. Cooke’s figure shows 22 cusps.
Dead. Durban (Sowerby, Smith); Natal (Fulton, and S. Afr. Mus.);
Inhambane and Mozambique Island (Braga).
Living. 28° 28’ S. 32° 25’ E. (off Cape St. Lucia), 27 metres (s.s. Africana).
Distribution. Red Sea, Zanzibar, Querimba Is., Mauritius, Réunion,
Seychelles, Madagascar, Indo-Pacific.
Remarks. The relatively small protoconch distinguishes this species from
heynemanni, apart from other characters. Smith’s mention of the small proto-
conch indicates that his two specimens were really young trapezium.
A Natal specimen, 53 x 24 mm. (S. Afr. Mus.) might certainly be regarded
as heynemanni because it is unicolorous without any trace of the dark spiral lines
so characteristic of trapezium; but it has the small protoconch (and narrower
ist and 2nd whorls) of trapezium.
The living specimen, 153 mm. long, dredged by the s.s. Africana has
indications of double lines only on the latest part of the outer lip, and only
feeble colourless plicae within; the base is quite smooth, without costae or
lirae; the protoconch is broken off, but was probably small.
That the characteristic dark spiral lines are not always developed in
juveniles and half-grown trapezium is shown by Smith’s (size not given), Fulton’s
69 mm., and the 8. Afr. Mus. 53 mm. examples. In addition to the one Natal
specimen, 5. Afr. Mus. also has examples from extra-African localities. Even
the 153 mm. Africana specimen was only just beginning to develop these dark
lines.
Fulton’s reasons for separating his strebeli from heynemanni were its more
slender shape and absence of well-marked spiral lirae; Smith also mentioned
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 79
these characters. But neither Smith nor Fulton mentioned the size of the proto-
conch, which, according to Fulton’s photographic figure, was small; nor did
they compare their specimens with trapezium.
A further difference between trapezium and heynemanni is: the former
develops six whorls against the latter’s four, in half-grown specimens of
approximately equal size.
Fasciolaria heynemanni Dnkr.
Fig. 18(d), 19(d)
1876. Kobelt in Kiister. Conch. Cab., p. 139, pl. 28, fig. 5.
1903. Von Martens. D. Tiefsee Exp., vii, p. 30.
1g11. Strebel. loc. cit., p. 28, pl. 5, figs. 27, 28; and p. 31, pl. 6, fig. 29 (scholvienc) ; and p. 33.
pl. 6, figs. 32, 32 a, b (dunkeri).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 46, pl. 4, figs. 3, 3a, 3b (alfredensis).
1923. Odhner. Géteb. K. Vet. Handl., xxvi, p. 6 (alfredensis).
1932. Turton. Mar. Sh. Pt. Alfred, p. 49; and p. 49, pl. x1, no. 362 (juv. dunkeri, non Strebel) ;
and p. 49 (scholviert, typ. err.); and p. 49 (alfredensis).
Not: Smith. Proc. Mal. Soc., v, 368 (= trapezium).
Spire 14-2 times in aperture. Profile of whorls with angular shoulder.
Protoconch 14 whorls, large, diam. 3°5—4°5, alt. 3-4 mm., smooth, last half
whorl with about 6 axial ribs, from suture to suture, continued on Ist postnatal
whorl, but restricted to the periphery and forming knobs, 5-6 spiral lirae above
the knobs, increasing to 8 or more on later whorls. Postnatal whorls 6, with
peripheral knobs varying in size, and in number from 9-14. Spiral lirae over
whole whorl, including the knobs; on base 8-10 more or less conspicuous very
flat costae, each traversed by about 3—5 lirae; usually only one, but sometimes
2 (scholuient) of these costae appear between the knobs and the suture of the
following whorl. On the rostrum the spiral lirae become almost axial in
direction (parallel with the canal), distinct in examples with straight rostrum
but obscured by the growth-lines in those with costate rostrum. Parietal callus
present. Columella with 3 pleats. Canal long, narrow, straight or slightly
sigmoid, especially when left side of rostrum is thickened and costate. Outer
lip slightly flattened in the middle and incurved below, not plicate internally,
in some examples with denticulate edge. 203 (protoconch and tip of canal
broken) xX 80 mm.; slenderest specimen 160 X 55 mm.
Operculum 62 X 33 in 203 mm. shell.
Pale buff or white, unicolorous, periostracum usually chestnut-brown,
but sometimes olivaceous golden-brown (Cape St. Blaize example) or golden-
brown (False Bay example); protoconch white; operculum dark brown.
Radula with c. 230-280 rows, lateral plate with 15 cusps on one side,
16 on the other, in another specimen 17 and 18 (excluding the tiny internal
denticle) ; middle cusp of the central plate asymmetrically bifid in one specimen.
Dead. Natal (Kobelt); Port Elizabeth, Port Alfred (Sowerby, Strebel,
Bartsch, Turton, $8. Afr. Mus.); Elim (i.e. Bredasdorp coast, Gape Agulhas
area) (Strebel: dunkeri); ‘Cape’ (Strebel: scholuvieni); off Glendower Beacon
.
80 ANNALS OF THE SOUTH AFRICAN MUSEUM
(Port Alfred area), 66 fathoms (S. Afr. Mus. P.F. coll.). Delagoa Bay S. Afr.
Mus. coll. K.H.B. 1912).
Living. Between Plettenberg Bay and St. Francis Bay, 100 metres (von
Martens); off Cape Infanta, 34-40 fathoms (Odhner) (also dead examples) ;
off Kowie, 40-43 fathoms; Algoa Bay, 40 fathoms; off Cape St. Blaize,
46 fathoms; False Bay (S. Afr. Mus. P.F. coll.). Cape Agulhas, littoral
CURE FE.
Remarks. A variable species, as was recognized by Strebel. He figured the
‘deep-water’ and the ‘coastal’ forms. The former has a straight non-costate —
rostrum, the latter a costate rostrum. But the present series shows that this
difference is not due to habitat, because both were obtained together in one
haul off the Kowie (P.F.); nor does it seem to be sexual because out of 5
animals 2 gg and 1 @ are non-costate, 2 99 costate.
This costate thickening of the rostrum seems to appear only in specimens
approximately 100 mm. in length upwards.
At the Kowie locality plump and slender examples were taken in the same
haul: width approx. 2-2—2-8 in the length, spire varying from 14, 132, to 2 times
in aperture (incl. canal), knobs g (in smallest specimen 58 mm. long) to 14 (the
2 largest examples 203 and 187 mm. had 11 and 13 knobs resp.).
The peripheral knobs are usually in the middle of the whorl, but may be
lower (more anterior); in dunkeri they adjoin the suture, except on the last
preserved whorl.
In some juveniles 2 subequal series of knobs are developed (cf. dunker?).
It is difficult to decide where the protoconch ends and the ist postnatal
whorl begins. The apex of the subglobular protoconch (14 whorls) is white,
the 1st postnatal whorl brown, but there is no sharp division between the two.
The axial ribs begin on the last part of the white region and are continued,
without any obvious interruption of growth, on the brown Ist whorl. One
specimen of protoconch is very large: diam. and alt. both 4:5 mm.
The curve of the outer lip helps to distinguish this species from lugubris,
but not from trapezium.
Strebel suggested that scholuient might possibly be a large form of the
dwarf heynemanni; but the present material negatives this; the differences are
merely individual.
Bartsch’s alfredensis was based on a worn slender specimen. Although the
plump, short-spired forms with strong knobs look different from the slender,
high-spired forms with weaker knobs, the present S. Afr. Mus. material, meagre
as it is, exhibits transitional forms.
F. dunkert, based on one young specimen (41:5 X 19:3 mm.), I regard as
an aberration. In some young examples of typical heynemanni the knobs on the
early whorls are very close to the suture of the following whorl; and in some
very young specimens a double row of feeble knobs occurs; but I have seen no
specimen exactly corresponding with dunkeri, i.e. with a double peripheral
shoulder adjoining the suture.
~
2)
2 i
5 ar =
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 81
In spite of Kobelt’s record from Natal, the Delagoa Bay locality is remark-
able. I cannot doubt the provenance because I myself found the specimen.
One would expect to find trapezium there, but the specimen is a typical heyne-
mann without a single one of the features distinguishing the Indo-Pacific
species. It retains the periostracum, and is thus unlikely to have been carried
very far by currents; moreover the Mozambique current sets in the wrong
direction to have carried this shell from Natal or the Port Alfred area.
Gen. Latrirus Mont.
1810. Montford. Conch. Syst., u, p. 530.
1840. Swainson. Treat. Malac., xxviii, p. 304 (Plicatella).
1891. Melvill. Mem. Manch. Philos. Soc., n. ser. 4, vol. 4, pp. 365-411 (Latirus + Peristernia).
1911. id. 7. Conch., xiii, p. 164 (Latirus + Peristernia).
Shell ovate or fusiform, usually ribbed. Canal long or short, columella
usually with pleats, aperture sometimes internally plicate. Operculum ovate,
apex curved inwards, nucleus apical, internal surface as in Fasciolaria (? all
species).
Radula central plate tricuspid, lateral plate with several cusps without
intervening smaller denticles.
The South African species fall into two groups:
Whorls with peripheral knobs—
abnormis, subcontractus.
Whorls with axial ribs and spiral irae—
polygonus, clausicaudatus, bairstowr, rousi, alboapicatus, turritus.
No living examples of bazrstowi Sow. 1886 or roust Sow. 1886 have yet been
found. The only South African record of turritus (Gmelin) is a dead shell from
Durban (Sowerby 1897); the species occurs at Mauritius, Seychelles, Ceylon,
etc.
Latirus abnormis Sow.
1894. Sowerby. 7. Conch., vii, p. 6.
1897. id. Append. Mar. Sh. S. Afr., p. 8, pl. 6, fig. 7.
1902. id. Mar. Invest. S. Afr., ii, p. 96, pl. 2, fig. 1 (émbricatus).
Fgoz. id. -ibid., p. 2277.
tgit. Melvill. 7. Conch., xiii, p. 165.
Aperture (incl. canal) a little longer than spire. Protoconch 2 whorls,
diam. 1-7, alt. 1-3-1-5 mm., smooth, junction with Ist postnatal whorl distinct.
Postnatal whorls 7, profile concave above the periphery, in some specimens
rather strongly so, almost forming a subsutural groove. Axial ribs 10 on Ist
whorl, well marked on periphery but feeble above and below, decreasing to
7-8 peripheral knobs on later whorls; crossed by 4—5 spiral lirae on 1st whorl,
increasing by interpolation on later whorls, but evanescent on 6th and 7th
whorls; 2 of the peripheral lirae on whorls 2—5 stronger than the others, making
the knobs subcarinate, but obsolete on knobs on 6th and 7th whorls; spiral
“'
vt
@
%
82 ANNALS OF THE SOUTH AFRICAN MUSEUM
lirae continued on base, 2 of them in upper third (at posterior end of aperture)
stronger than the others and forming a series of feeble subcarinate knobs cor-
responding in position with the peripheral knobs. Parietal callus weak;
columella without pleats. Canal not abruptly separated from rest of aperture,
which is posteriorly more or less indented; columella glaze in large specimens
discrete from rostrum forming a narrow umbilicus; outer lip not plicate within.
Periostracum thin, fibrous, imbricate-scaly. 72 X 29 mm. (S. Afr. Mus.);
Brit. Mus. specimen (when perfect) probably 75 mm. long (Smith).
Operculum ovate, gently curved, 13 X 6:5 mm. in 49 mm. shell, internal
surface as in Fasciolaria.
Ochraceous salmon, or orange-brown, periostracum amber-brown,
operculum brown.
Living. Off Durnford Point (Zululand), 13 fathoms (S. Afr. Mus. P.F.
coll.).
Dead. Natal (probably from fish stomachs) (Smith); off Tugela River,
46 fathoms (Sowerby), and 14 fathoms (S. Afr. Mus. P.F. coll.).
Latirus subcontractus (Sow.)
1902. Sowerby. Mar. Invest. S. Afr., li, p. 97, pl. 2, fig. 2 (Fusus s.).
1932. Tomlin. Ann. S. Ajy. Mus., xxx, p. 158, fig. 2 (mosselensis).
Aperture a trifle longer than spire. Protoconch 2 whorls, diam. 1:3, alt.
1 mm. smooth, beginning of Ist postnatal whorl distinct (type of mosselensis).
Postnatal whorls 8; 9 (? 10) (subcontractus) or 8 (mosselensis) axial ribs on each
whorl, from suture to suture on 1st-3rd whorls, but from 4th whorl restricted
to the periphery; crossed on 1st whorl by 5 spiral lirae, increasing on following
whorls, but from about the 5th whorl evanescent, except the peripheral lira
which persists on the knobs (not in the intervals) making these subcarinate; on
base a second series of weaker subcarinate knobs, and a few weak lirae on
rostrum. Parietal callus weak, columella with 2 obscure pleats (mosselensis) ;
canal rather abruptly demarcated from rest of aperture, narrow (subcontractus) .
A narrow umbilicus (mosselensis) ; outer lip not plicate internally. Periostracum
thin, smooth. 40 * 18 mm. (subcontractus); 53°5 X 23 mm. and 60 X 25 mm.
(type and figure of paratype of mosselensis).
Operculum ovate, gently curved (Tomlin).
Pale pinkish, periostracum yellowish-brown (mosselensis type).
Dead: off Cape Natal (Durban), 200 fathoms (Sowerby).
Living: Mossel Bay, 27 fathoms (Tomlin) (both S. Afr. Mus. P.F. coll.).
Remarks. Has a strong resemblance to a Fasciolaria, cf. heynemanni or
trapezium. ;
Type of subcontractus ? in British Museum. Tomlin figured the paratype
(? in coll. Tomlin) with operculum, and the back view of the type of mosselensts.
The latter is in S. Afr. Mus. without operculum. The radula was not described.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 83
Tomlin compared his species with L. armatus A. Adams 1854 (see 1886.
Challenger Rep., xv, pl. 13, fig. 1, Fasciolaria armata), but made no reference to
_ the truly remarkable resemblance to Sowerby’s species.
Except that subcontractus appears (from Sowerby’s figure) to have one
(? 2) more axial ribs (knobs) than mosselensis, and that the latter is rmmate and
has a slightly curved columella, there are no differences between Sowerby’s
figure and the type of mosselensis. The lower part of the canal and outer lip
are broken in the latter, which may account for the umbilicus being visible.
In spite of the distance apart of the two localities, and the difference in
depth, one cannot accept more than the one species.
Latirus polygonus (Linn.)
1816. Lamarck. Tabl. Encyclo., pl. 423, fig. 1 (Fusus p.).
1859. Chenu. Man. Conchyl., i, fig. 908.
1903. Smith. Proc. Mal. Soc., v, p. 369 (var.).
The only South African record is from ‘Durban, deep water’ [probably
from fish stomach] (Smith). The species occurs at Mauritius, and other
localities in the Indian Ocean. Specimens in S. Afr. Mus. from ee (coll.
Rawson W. Rawson).
The coloration appears to be distinctive: buff or ochraceous, with brown
axial ribs divided into oblong patches by the pale spiral lirae.
Latirus clausicaudatus (Hinds)
1844. Hinds. ool. Voy. Sulphur. Moll., p. 13, pl. 1, figs. 10, 11 (Fusus c.).
1892. Sowerby. Mar. Sh. S. Afr., p. 3 (Fusus c.).
Elongate-fusiform, turreted, aperture (incl. canal) 14—1} times spire.
Profile of whorls evenly convex. Protoconch 24 whorls, diam. and alt. 2 mm.,
smooth, junction with rst postnatal whorl! distinct. Postnatal whorls 7; 7 axial
ribs on Ist whorl, increasing to 8 or g on last whorl, from suture to suture but
becoming weaker above periphery on later whorls, and evanescent on lower
two-thirds of base; sometimes indistinct on last part of 7th whorl, but
reappearing as a well-marked rib on outer lip; crossed by spiral rae, 7-8 on
Ist whorl increasing to 16-20 on later whorls, 35-40 additional on base.
Parietal callus bluntly dentiform; columella with 2 pleats, the anterior one
distinct, the other obscure; canal abruptly marked off from rest of aperture,
very long, nearly twice the rest of aperture, nearly closed throughout its length
by the discrete edge of the columella glaze, no umbilicus. Aperture internally
without plicae except one at base of canal opposite the columella pleat. Perio-
stracum thin, smooth. 51 X 15°5 mm., smallest specimen in 8S. Afr. Mus.
31 mm. long. Hinds’s figure 58 xX 16-5 mm.
Operculum oval, apex incurved, 7 X 3°5 mm. in 45 mm. shell.
84 ANNALS OF THE SOUTH AFRICAN MUSEUM
White or pale buff, periostracum pale greyish brown, operculum dark
brown. :
Dead: Agulhas Bank, 50-60 fathoms (Hinds). Off Cape Natal (Durban),
54 fathoms; off Cape Morgan, 77 fathoms; off Nahoon Point (East London),
45 fathoms; off Hood Point (East London area), 49 fathoms; off Nanquas
Peak (eastern part of Algoa Bay), 63 fathoms; Algoa Bay, 37 fathoms; (S. Afr.
Mus. P.F. coll.).
Living: off Riet Point (east of Algoa Bay), 23 fathoms (S. Afr. Mus.
PE, jcoll:):
Remarks. These specimens are clearly referable to Hinds’s species, which
Sowerby (1892, also 1903. Mar. Invest. S. Afr., 1, p.. 97) said was represented.
by the unique type in the British Museum.
In one specimen the protoconch and 3 whorls are slightly curved to the
right, and in another the protoconch is curved to the left as in Hinds’s figure.
The animal of the only specimen taken alive was not preserved.
Latirus alboapicatus Smith
1902. Smith. 7. Conch., x, p. 250, pl. 4, fig. 5.
1906. Smith. Ann. Natal Mus., i, p. 34, pl. 7, fig. 7 (burnupt).
1911. Melvill. 7. Conch., xii, p. 166 and p. 168 (burnupt).
1931. Tomlin. Ann. Natai Mus., vi, p. 433.
Fusiform. Profile of whorls slightly angularly convex (fig. of alboapicatus),
slightly concave above, then gently convex (burnupi). Protoconch 2 whorls,
diam. and alt. ¢. 1-5 mm., smooth. Postnatal whorls 6; axial ribs c. 12 (albo-
apicatus) on 1st whorl (in burnufi corroded), 8 on 2nd and following whorls, from
suture to suture, but more prominent on the periphery, and evanescent on
base; crossed by spiral lirae (number not stated in alboapicatus) 5 on and whorl
increasing to 8-10 on last whorl, the first one or first 2 or 3 below suture granu-
lose, 9-11 additional on base, of which the 4th or 5th is stronger than the
others, covered posteriorly by the parietal callus and forming a small denticle
on outer lip; columella with 3 weak pleats, canal straight (slightly recurved in
alboapicatus), distinctly but not abruptly marked off from rest of aperture,
umbilicus slight or absent, outer lip sometimes plicate internally. Periostracum
thin, smooth. 28 X 11°5-12 mm.
Operculum (burnupi) ovate, apex incurved, 6 X 3 mm. in 28 mm. shell.
Rufous with white apex, and a pale band below centre of body whorl,
aperture rufescent within (alboapicatus); white with brown periostracum, the
strong lira on base showing as a pale line, aperture rosy or purplish within,
operculum dark brown (burnup) (see Remarks).
Dead: (alboaficatus) Durban (Tomlin). Living: (burnupi) Port Shepstone
(Natal) (Smith, also S. Afr. Mus. Ross-Frames coll. ex Burnup).
Remarks. Although Smith must have had his alboapicatus for comparison
when he described burnupi, and Melvill accepted both species, I strongly suspect
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 85
that the two are conspecific. As I have seen no specimens of the former species,
the description is based mainly on specimens of burnupi collected in the type
locality by Burnup.
Like Smith’s specimens, all these are more or less corroded at the apex,
so that the true size of the protoconch or the sculpture of the 1st whorl cannot
be determined.
Tomlin (1931) records (fide Burnup) a specimen of alboapicatus 44 (45) X
16-5 mm. (!), of which Burnup said the ribs and growth-lines were paler than
the intervening spaces, and the spiral grooves much darker than the lirae. This
applies also to Burnup’s specimens of burnupi in S. Afr. Mus.
Gen. PERISTERNIA Morch
1852. Morch. Cat. Conch. Yoldi., i, p. 99.
1891. Melvill. Mem. Manch. Philos. Soc. (4), iv, p. 365 (Latirus part).
1911. id. 7. Conch., xiii, p. 164 (Latirus part).
1935. Yen. Notes Malac. Chinoise, i (2), p. 41 (Peristerina emend.).
Shell broadly fusiform, more or less distinctly axially ribbed and spirally
lirate, the lirae often scabrous or squamose; not or only slightly umbilicate,
canal moderate, slightly recurved, columella usually with 2 pleats, aperture
internally plicate. Operculum as in Latzrus.
Radula: central plate piriform, narrowed in front, with 3 feeble cusps, and
lateral plate with denticles between the main cusps.
Included conchologically in Latirus, by Melvill, but can be distinguished
by the radula.
Peristernia leucothea Melv.
Fig. 19(e)
1891. Melvill. loc. cit., p. 399, pl. 2, fig. 15.
1900. Sowerby. Proc. Mal. Soc., iv, p. 1, pl. 1, fig. 2 (Euthria eburnea).
? 1932. Turton. Mar. Sh. Pt. Alfred, p. 54, pl. 12, no. 400 (Euthria ordinaria).
1937. Peile. 7. Conch., xx, p. 300, fig. 1 (radula).
Length about twice breadth. Aperture subequal to spire. Protoconch 14
whorls, diam. 0:8, alt. 0-75 mm., smooth, 2 riblets close together before junction
with Ist postnatal whorl. Postnatal whorls 7; axial ribs g-10 on 1st whorl,
increasing to 11-12 on last whorl, from suture to suture but more prominent
peripherally, evanescent on base; crossed by spiral lirae, 3 on Ist and 2nd
whorls, 4 on 3rd, 5-6 on 4th, increasing to 10-11 on last whorl, often with
intermediaries, 10-12 additional on base, also with intermediaries, usually one
or two at about the middle of base stronger than the others; fine close-set
growth-lines producing punctae in the sulci between the lirae. Parietal callus
present, columella with 3 pleats but usually only 2 or one distinct, forming a
short keel at beginning of the short canal. Outer lip internally plicate, the 1st
86 ANNALS OF THE SOUTH AFRICAN MUSEUM
(posterior) and last (opposite columella pleat) plicae larger than the others,
more or less dentiform. Columella glaze sometimes rimate anteriorly forming
a feeble umbilicus. Periostracum very thin. 25 X 12 mm.
Operculum ovate, apex incurved, 4°5 < 2°5 mm. in 22 mm. shell, internal
surface as in Fasciolaria.
Creamy-white or pale buff, unicolorous; or orange-brown with markings,
the colour when present is mostly around the suture, between the ribs, and in
one or two bands on base, aperture internally white or pale brown, or pale
violaceous, operculum chestnut-brown, periostracum yellowish-brown.
Radula with 250-270 rows, lateral plates with g-11 (12) cusps and
denticles, not always symmetrical and the sequence of cusps and denticles
varying from one part of the radula to another.
Dead: Port Natal (Durban) (Melvill), Isipingo and Umkomaas (Natal)
(Smith), Tongaat (Natal) (S. Afr. Mus.); Pondoland (Sowerby: eburnea) ;
Port Alfred (Turton: ordinaria).
Living: off Durnford Point (Zululand) 13 fathoms (S. Afr. Mus. P.F.
coll.); Scottburgh (Natal) littoral (S. Afr. Mus. coll. K.H.B.); Umpangazi,
Umbhlali, Durban, Umtwalumi, Port Edward and Port St. Johns (U.C.T.).
Remarks. The apex is usually corroded in littoral specimens; only one of
the numerous Scottburgh specimens had a complete unworn protoconch.
Appears to be an ‘albino’ form of nassatula, as there is no conchological
difference between the two. Of the specimens I have seen, those most strongly
marked with orange-brown come from Umblati, Tongaat, and Durban.
Others from the last locality are uniform white or buff; one of the Umpangazi "
shells has a pale violaceous aperture. For the present retained separate from
nassatula.
One specimen with an aberrant operculum: oval, nucleus intramarginal
in apical third of length.
A series of water-worn specimens (S. Afr. Mus. locality?) is interesting.
The effect of wear is to broaden the ribs and reduce the intervening grooves in —
which the spiral lirae, or their intervening sulci, persist, though much reduced.
Even when completely worn away at the upper part of a whorl, they usually
persist in the lower part adjoining the suture of the following whorl. The final
result—a perfectly smooth shell—appears to be represented by Turton’s
ordinaria from Port Alfred, the locality farthest removed from the habitat of
the living animal.
Even badly worn specimens, however, are too broad to be mistaken for
Suscotincta.
Fic. 19. ST
Central and lateral radula plates of (a), (b) Fasciolaria lugubris Rve. from juvenile from egg-
capsule, and from 85 mm. shell; (c) F. rutila Watson; (d) F. heynemanni Dnkr.; (e) Peristernia
leucothea Melv.; (f) P. fuscotincta (Sow.); (g) Fusus verruculatus Lam.; (h) F. faurei n. sp.;
(1) F. rubrolineatus Sow., two variants of lateral plates; (j) F. colus Linn., half-grown and adult,
the latter with abnormal 4-cuspid central plate; (k) F. africanae n. sp.
88 ANNALS OF THE SOUTH AFRICAN MUSEUM
P. incarnata Desh., recorded from Natal (Sowerby 1892) (occurs also at
Mauritius, Red Sea, and Indo-Pacific) differs from Jleucothea in having fewer
spiral lirae on last whorl (6-7 in Philippine specimens in S. Afr. Mus.),
and in coloration, which is yellow or orange with brown intervals between
the ribs.
Peristerna nassatula (Lam.)
1859. Chenu. Man. Conchyl., i, fig. gto.
1880. Von Martens. Mauritius G Seychellen, p. 246 (Plicatella n.).
The description given for leucothea will apply to this species which is,
however, more brightly coloured.
Cream, upper half of whorls and the grooves between the ribs brown,
shading off into orange-brown, base with a pale spiral band at level of top of
aperture, followed by a dark brown band and then another pale band, rostrum
orange-brown, aperture violaceous.
Radula (one specimen from Delagoa Bay examined) incomplete but with
at least 180 rows, lateral plate with (8) 9-10 cusps and denticles, varying in
size and sequence as in leucothea.
Natal (Krauss, ? dead). Delagoa Bay, living (U.W.).
Distribution. Mauritius, Réunion, Seychelles, East Indies.
Peristernia fuscotincta (Sow.)
Fig. 19(f)
1886. Sowerby. 7. Conch., v, p. 2 (Euthria f.).
1689, id. “abid, wi, pli, fies 18.
1892. id. Mar. Sh. S. Afr., p. 4, pl. 1, fig. 13 (Huthria f.).
1938. Peile. Proc. Mal. Soc., xxiii, p. 99, fig. 36 (radula).
1947. Stephenson. Ann. Natal Mus., xi, pp. 271, 273 (Cominella f.).
Length distinctly more than twice breadth. Aperture shorter than spire.
Protoconch 2 whorls (all specimens worn). Postnatal whorls 5; faint indications
of weak ribs on upper 2 or 3 whorls; all whorls with spiral grooves, 4 on 2nd
whorl increasing to 10-11 on last whorl, usually in pairs, and punctate where
crossed by the growth-lines, g-10 additional on base, some in pairs. Parietal
callus present, columella with 3 pleats, but only one or two distinct, forming a
short keel at beginning of the short canal. Outer lip internally plicate, 1st and
last plicae larger than the others. Sometimes a feeble umbilicus. 20 x 8 mm.
Operculum oval, apex incurved.
White with irregular brown markings, either as patches or axial flames,
usually a more or less continuous brown band. below periphery.
Radula with 140-160 (Peile: 167) rows, lateral plate usually with 6 major
cusps with intervening denticles (1 or 2); the arrangement varies in successive
rows, and is not always symmetrical on the two sides (cf. leucothea).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 89
P.. fuscotincta: dead; Port Elizabeth, Port Alfred (Sowerby, Bartsch,
Turton); Port Shepstone (Natal) (S. Afr. Mus. coll. Burnup).
Living: Port St. Johns to Richmond (Alexandria Division) (Stephenson) ;
East London to Kleinmond (Bathurst Division) (U.C.T.).
Remarks. In fuscotincta the spiral grooves were described as very obscure
(they certainly are in beach-worn specimens!).
Some of the beach-worn specimens in S. Afr. Mus. are almost wholly
white, but even so the subperipheral band and a few brown marks are just
visible.
Stephenson (1947 p. 271 footnote) transferred this species to Cominella
(Afrocominella) seemingly at Tomlin’s suggestion—a clear case where con-
chological guessing proved wrong —although Peile’s examination of the radula
had already (1938) put the species in its correct genus.
Gen. Fusus Brug.
1789. Bruguiére. LEncycl. Meth. (1), xv.
Shell more or less elongate-fusiform spire often high. Canal moderately
or very long; columella without pleats but sometimes rimate.
Operculum oval or piriform apex blunt more or less incurved or sharply
pointed, nucleus apical, internal surface with marginal thickening as in
Fasciolaria.
Radula, central plate subtriangular, narrowed in front, tricuspid, lateral
plate with several cusps, without intermediate denticles.
Remarks. F. radialis Watson has proved, not unexpectedly, to be a
Columbarium (p. 234), and F. speratus a Tritonalia (p. 215).
Fusus africanus (Sow.)
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 1, pl. 6, fig. 19.
1903. Smith. Proc. Mal. Soc., v, p. 368, pl. 15, fig. 19.
1932. Turton. Mar. Sh. Pt. Alfred, p. 50, pl. xi, no. 370 (kowzensis).
Piriform. Aperture (excl. canal) a little longer than spire (incl. canal:
24-24 times spire). Profile of early whorls nearly straight, of later whorls
angular. Protoconch 2 whorls, smooth (but no perfect specimen seen). Post-
natal whorls 6; on first 3 whorls 10-11 very obscure axial ribs, reduced on
following whorls to peripheral rounded knobs, which become stronger on last
2 whorls: on 4th whorl near suture, on 5th and 6th nearer middle of whorl;
crossed by spiral lirae, 5 on 1st whorl, increasing to 8-g (10) on 4th whorl but
thereafter evanescent; on base 12-15 low blunt costae, evanescent towards end
of rostrum. A blunt parietal callus, columella curved, glaze discrete forming
an umbilical rimation; canal long, straight, narrow, distinctly marked off from
go ANNALS OF THE SOUTH AFRICAN MUSEUM
rest of aperture; outer lip not plicate within. Periostracum thin. 104 X 52mm.
(Turton); 79 (protoconch missing) xX 38 mm. (S. Afr. Mus.).
Operculum and radula unknown.
Creamy, buff, pale orange-brown, with darker marks between the knobs
and sometimes axial flames, grooves between costae on base orange-brown,
periostracum brown.
Port Elizabeth (Sowerby); Port Alfred (Turton: kowiensis); off Durban
[from fish stomachs] (Smith, also S. Afr. Mus.).
Remarks. Only dead specimens known. Except for Sowerby’s original
young specimens, and Turton’s specimen, this species has only been obtained
from fish stomachs in Natal waters. Not taken by the Pieter Faure.
Fusus verruculatus Lam.
Figs. 19(g), 20(a)
1816. Lamarck. Tabl. Encycl. Meth., p. 429, fig. 7, and Liste, p. 7 (ocelliferus, name and
figure only).
1822. Id. Anim. sans Vert., vii, p. 129.
1870. H. Adams. Proc. Zool. Soc. Lond., p. 110, text-fig. (ventricosus, non Gray).
1876. Kobelt. Conch. Cab., p. 152, pl. 47, fig. 3 (adamsit).
1886. Watson. Challenger Rep., xv, p. 195 (references).
1892. Sowerby. Mar. Sh. S. Afr., p. 3 (robustior).
1925. Thiele. D. Tiefsee Exp., xvii, p. 184, pl. 32 (20), fig. 19 (juv. referred with ? to
capensis Thiele).
1932. Turton. Mar. Sh. Pt. Alfred, p. 50 (ocelliferus and robustior), pl. xi, no. 369 (robustior
juv.).
Aperture (incl. canal) 14 to nearly 14 times spire. Profile of early whorls
slightly angular, of later whorls angular (with knobs) (verruculatus), convex
(adamsi). Protoconch 24 whorls, diam. and alt. 1-5-1:75 mm. (rarely perfect
except in juv.), smooth but with some irregular plicae before the abrupt
junction with 1st postnatal whorl. Postnatal whorls 8; axial ribs 9-10 on Ist
whorl, increasing to 11, usually from suture to suture on first 3 whorls, but
thereafter only at periphery where they become blunt, more or less complanate
knobs, 11-15 in number, continuing on to 8th whorl forming a prominent
shoulder in typical verruculatus, but petering out on 7th or 8th whorl in adamsiz ;
crossed by spiral lirae, 3 (4) on 1st whorl, increasing to 5 (6) on 2nd and grd
whorls, end and 3rd lirae strongest, thereafter the 3rd strongest, on the periphery
and carrying knobs, i.e. 2 lirae above and 2 (sometimes 3) below the peripheral
keel; 15-20 additional lirae on base, with intermediaries; also over the whole
whorl fine spiral striae. Parietal callus not strong, sometimes in addition with
3-4. (up to 6) plicae; columella curved, rimate in half-grown to adult examples,
forming a deep but narrow umbilicus. Canal nearly straight in juv., usually
more strongly curved in older examples, subequal to and not sharply marked
off from rest of aperture. Outer lip at some stages of growth plicate within.
Periostracum thin, fibrous and imbricate, especially near the suture, fimbriate.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA QI
(verruculatus) 150 (protoconch and tip of rostrum broken, say: 153) X 70 mm.;
119g X 45 mm.; (adamsiz) 133 X 48 mm.; 125 X 50 mm.
Operculum oval, apex incurved, 35 < 18 mm. in 125 mm. shell.
Cream or buff, the peripheral knobs usually reddish-brown, sometimes
also indications of orange flames, periostracum greyish or yellowish-brown,
operculum horny or reddish-brown; juveniles seem to. be more brightly
coloured (at least in some beach-worn examples the coloration shows better),
with darker knobs and flames.
Animal bright red with minute white specks (K.H.B.).
Radula in 7 mm. shell with 110 rows, lateral plate with 7 cusps, in 23 mm.
shell resp. 165 and 8, in 25 mm. shell resp. 170 and 8, in 30 mm. shell resp. 205
and 9, in 38 mm. shell resp. 215 and g, in 62 mm. shell resp. 230 and 11-12, in
114 mm. shell resp, 285 and 12, in 150 mm. shell resp. 325 and 13 (the tiny
denticle at inner end excluded in all counts).
Dead and beach-worn specimens recorded from Port St. Johns, Port
Alfred, Port Elizabeth, Agulhas Bank, Still Bay, False Bay (Sowerby, Adams,
Bartsch, Turton and 8. Afr. Mus.).
5° 16’ S. 22° 26’ E., 155 metres (Thiele, juv.).
Living: Simon’s Bay (False Bay), 15-20 fathoms (Watson); Algoa Bay
and Agulhas Bank to mouth of False Bay, 10-66 fathoms (S. Afr. Mus. P.F.
coll.). Sea Point (Cape Town), low tide (S. Afr. Mus.), Knysna, low tide
(S. Afr. Mus. P.F. coll.). Both the latter adamsii form). 33° S. 28° 11’ E. (off
East London), 31 fathoms (verruculatus form) (U.Q.T.).
Saldanha Bay, 10-14 fathoms (S. Afr. Mus. P.F. coll.).
34° 35'S. 19° 14’ E., 66 metres; False Bay, 3-24 metres; west coast of
Cape Peninsula, intertidal; Langebaan (Saldanha Bay); off Lambert’s Bay,
66 metres (U.C.T.). )
Remarks. The East London locality bridges the gap between Port St.
Johns and Port Alfred.
The verruculatus and adamsi forms are not restricted to separate areas.
Juveniles from 5 mm. long (protoconch plus 1st whorl) examined.
While the early whorls show little variation, the later whorls show marked
dimorphism: the typical verruculatus with strong shoulder knobs, and adamsii
with evanescent knobs and evenly convex whorls. The institution of adamsii
as a distinct species is not surprising when only the extreme forms were available.
But they are connected by transitional forms.
Plump and slender examples occur in both verruculatus and adamsii, though
the latter in general is the more slender. The most slender specimen I have
seen is one (adamsi) taken in False Bay measuring 118 x 40 mm.; it is not
scalariform but the spire is elongated to such an extent that its length equals
the length of aperture (incl. canal).
In the early whorls the axial ribs are usually well developed (see Thiele’s
figure), and the bicarinate periphery on the 2nd and 3rd (sometimes also but
less conspicuous on 4th) whorls is very characteristic.
Q2 ANNALS OF THE SOUTH AFRICAN MUSEUM
One specimen (S. Afr. Mus. no. A4661, off Cape St. Blaize) 82 mm. long,
and a juvenile (locality ?) 20 mm. long, have unusually large protoconchs:
diam. and alt. almost 2:3 mm.
Another specimen, also from off Cape St. Blaize, 147 x 57 mm., has
peripheral knobs extending on to the 8th whorl but the profile is convex, not
shouldered; and the canal is markedly sigmoid.
One specimen (S. Afr. Mus. no. A4662, off Cape St. Blaize) 137 (tip of
canal broken, probably 140-142 when perfect) x 66 mm., is subscalariform,
with strongly convex ventricose whorls, and deeply sunken sutures.
It is a question whether the name ocelliferus in Lamarck’s Liste des objets
(sometimes attributed to Bory, but see: Sherborn & Woodward. Proc. Zool.
Soc. Lond., 1893, p. 584) should be used for this species. The figure is recog-
nizable as a representation of this species, but is it adequate to i it
from other species?
Fusus colus Linn.
Figs. 19(j), 20(d)
1816. Lamarck. Tabl. Encycl., pl. 423, fig. 2, and Liste, p. 6 (longicauda).
1859. Chenu. Man. Conchyl., i, fig. 597.
1876. Kobelt. Conch. Cab., p. 146, pl. 30, fig. 3, pl. 47, fig. 1.
1942. Gravely. Bull. Madras Govt. Mus., V, 2, p. 62, fig. xi, i (longicauda).
1952. Satyamurti. ibid., n.s. I, 2 (6), p. 187 (longicauda).
Protoconch 24 whorls, diam. 1, alt. 1-3 mm., smooth, with a dozen or
more fine axial ribs in last half whorl, which is more or less sharply demarcated
from ist postnatal whorl.
Radula in 25 mm. shell with 80 rows, lateral plate with 8 cusps, in 40 mm.
shell resp. 140-170 and 8-9, in 97 mm. shell c. 220 and 11 (excl. the minute
inner denticle); central plate broader than long, narrowed in front, tricuspid
(in one specimen 4-cuspid).
Living: off Umhloti and Umvoti Rivers (Natal), 25-27 fathoms; off
Amatikulu River (Zululand), 24 fathoms (S. Afr. Mus. P.F. coll.). Delagoa
Bay (S. Afr. Mus. coll. K.H.B., and U.W.). Inhambane (U.C.T.).
Dead: off Tongaat and Umhlanga (Natal), 22-36 fathoms (S. Afr. Mus.
P.F. coll;
Remarks. F. toreuma (Martyn), recorded from Natal by Smith (1903), is
distinguished by the angular profile of the whorls. If the Natal specimen was
taken from a fish stomach, the species is probably living in South African
waters.
Fusus torulosus Lam.
1816. Lamarck. Tabl. Encycl. Meth., pl. 423, fig. 4, and Liste, p. 6.
Specimen in S. Afr. Mus., probably from Ceylon or Indian Ocean.
Aperture (incl. canal) longer than spire. Protoconch missing. Postnatal
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 93
whorls 10; profile convex with crenulations due to the spiral lirae. Axial ribs
5-6 on 1st and 2nd whorls, increasing to 10-11 on last, broad and rounded,
from suture to suture; crossed by sharp spiral lirae 4—5 on 1st whorl, increasing
to 10-11 on last, with an intermediary between the peripheral pair; c. 35
additional lirae on base and rostrum, some of them feebler than others. Growth-
lines distinct immediately below suture, more or less so between the lirae on
rest of whorl. Sutures deep. 91 (protoconch missing) X 24 mm.
A fragment of two half whorls exactly agreeing with the sculpture of the
above described specimen: off Cape Natal, 85 fathoms; off Umhloti River,
40 fathoms, 2 worn fragments; off O’Neil Peak (Zululand), 90 fathoms, one
worn fragment (S. Afr. Mus. P.F. coll.).
Fusus rubrolineatus Sow.
Fig. 19(2)
1870. Sowerby. Proc. Zool. Soc. Lond., p. 252.
1880. id. Thes. Conch., iv, p. 80, pl. 411, fig. 68.
1903. id. Mar. Invest. S. Afr., ii, p. 228.
1903. Von Martens. D. Tiefsee Exp., vii, p. 30.
1903. Thiele. ibid., p. 169, pl. 9 (4), fig. 60 (radula).
1915. Bartsch. Bull U.S. Nat. Mus., 91, p. 47.
Aperture (incl. canal) 1-1} times spire. Protoconch 24 whorls, diam.
and alt. 1:2 mm., smooth, with 4-6 axial ribs on last half whorl, junction with
Ist postnatal whorl distinct. Postnatal whorls 7; axial ribs 14-15 on 1st whorl,
increasing to 17-18 on last whorl, from suture to suture, but evanescent towards
suture on last whorl and on lower half of base, from about the 4th whorl slightly
curved. below the suture; fine close-set growth-lines; crossed by 3 spiral lirae
which appear alone on 1st whorl and continue as 3 main lirae on all following
whorls but with added intermediaries (e.g. on 6th whorl 6—7 between suture
and Ist main lira, 3 between Ist and 2nd, and between 2nd and 3rd, 3-4 below
grd lira); 15-20 additional lirae on base, those on upper half with inter-
mediaries; main lirae on the whorls and upper half of base forming horizontal
complanate nodules at intersections with axial ribs. No parietal callus, colu-
mella slightly curved, in some specimens with a slight swelling (scarcely a
pleat) at the bend. Canal shorter than, but not marked off from rest of aperture,
slightly curved. Periostracum thin, smooth. 38 x 13 mm., a plump specimen
32 X 13 mm.
Operculum broadly oval, apex rounded, slightly on outer side of median
line (von Martens said toward inner side), 6 X 4 mm. in 35 mm. shell, internal
surface as in Fasciolaria. -
Pale buff, main lirae on whorls and base, and also some of the more
prominent intermediaries orange-brown, forming spiral lines, usually con-
tinuous but often more intense on the axial ribs, producing an effect of series
of spots or axial flames.
Q4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Radula with 150-180 rows, central plate longer than wide, narrower in
front, with 3 rather strong cusps, lateral plate with 5-6 cusps (excl. inner
denticle), varying in size, and often asymmetrical.
Agulhas Bank (Sowerby 1870); 35° 16'S. 22° 26’ E., 155 metres (von
Martens); off Cape St. Blaize, 53 fathoms (Sowerby 1903).
Living and dead: Agulhas Bank, from approximately 22° E. to 274° E.,
and southwards to Brown’s Bank approx. 363°S., 63-124 fathoms (S. Afr.
Mus. 'P:F. coll:).
Remarks. Von Martens (loc. cit. 1903. p. 103, pl. 2, fig. 10) described
rufinodis from off Zanzibar and Sumatra, and compared (p. 104) it with rubro-
lineatus. But he seems to have compared his specimens with Sowerby’s figure
instead of with actual specimens of rubrolineatus which were available to him.
The differences are not very convincing and the two species are obviously
closely allied. F. rufinodis, however, has only 10 axial ribs on the 7th (pen-
ultimate) whorl and 11-12 on the 8th whorl; and the central plate of the radula
(Thiele, loc. cit., fig. 59) is nearly square.
An even more closely allied species is F. libratus Watson 1886 from Fiji
Islands, 315 fathoms.
Most of the Pieter Faure examples are, like the Valdivia specimens, covered
with an Alcyonarian.
Fusus faurei n. sp.
Figs. 19(h), 20(e)
Aperture (incl. canal) 14 times spire. Profile of whorls convex, slightly
biangulate. Protoconch 2 (24) whorls, diam. and alt. 2 mm., smooth (but all
specimens more or less corroded). Postnatal whorls 6; axial ribs 11-12 on Ist
whorl, increasing to ¢. 18-20 on last whorl (irregular and obscure on last half
whorl), low, rounded not prominent except at the 2 peripheral lirae, slightly
retractive below suture; crossed by 2 main peripheral costae from 1st whorl
onwards, with finer lirae above and below, on last whorl 6-7 above and 3-4
below the peripheral costae, varying in strength, the lowest one (next the
suture) the strongest, 2-3 between the 2 costae, 15-20 additional lirae on base,
usually an intermediary between each pair. No parietal callus. Columella
gently curved, no pleats, not rimate. Canal a little shorter than, and distinctly
but not sharply marked off from rest of aperture, straight or very slightly
reflexed at tip, open. Periostracum thin. 50 (protoconch and tip of canal
broken) X 22 mm.; 33 X 15 mm.
Operculum oval, apex blunt, incurved, 9 X 5 mm. in 38 mm. shell.
Creamy-white, periostracum grey, operculum amber-brown.
Radula with at least 180 rows, central plate narrowed in front, tricuspid,
lateral plate with 11 cusps (excl. inner denticle).
Living and dead: off Cape Point, 300-560 fathoms (S. Afr. Mus. A4581
(Type), Aq58e. PF. coll).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 95
Fusus bonae-spei n. sp.
Fig. 20(c), (f)
1925. Thiele. D. Tiefsee Exp., xvii, p. 183, pl. 32 (20), fig. 18 (capensis, non Dunker).
Not the juv., p. 184, fig. 19 = verruculatus.
Aperture (incl. canal) a little longer than spire. Profile of whorls convex
but weakly biangulate. Protoconch 24 whorls, when not corroded diam.
2 mm., alt. 2-5-3 mm., smooth, junction with 1st postnatal whorl abrupt.
Postnatal whorls 7; feeble axial ribs 8—g on Ist whorl, increasing to g-10, but
not traceable beyond 4th whorl; growth-lines distinct; crossed by spiral lirae,
on Ist whorl 3, the lower two stronger than the upper one, continued on
following whorls, the two stronger ones forming the periphery, with weaker
intermediaries producing a fine cancellate sculpture, 20-25 additional lirae on
base also with intermediaries. No parietal callus, columella slightly curved,
Fic. 20.
(a) Fusus verruculatus Lam., protoconch, with detail of one lira. (6) F. colus Linn. protoconch.
(c) F. bonae-spei n. sp. protoconch. (d) F. africanae n. sp., protoconch. (e) F. faurei n. sp., profile.
(f) F. bonae-spet n. sp., profile.
96 ANNALS OF THE SOUTH AFRICAN MUSEUM
canal 13-14 times, but not very sharply marked off from rest of aperture,
straight or slightly curved and reflexed, narrow. Outer lip not plicate inter-
nally. Periostracum thin, fibrous-fimbriate. 102 X 34 mm.
Operculum oval, apex incurved, 20 X 12 mm. in 102 mm. shell, internal
surface as in Fasciolaria.
Creamy-white or pale buff, periostracum yellowish-buff, operculum brown.
Dead: 34° 33’ S. 18° 2’ E., 318 metres (Thiele).
Living and dead: off Cape Point, 85-256 fathoms (S. Afr. Mus. A4622,
A4628-32, Type A4632. P.F. coll.).
Remarks. I have not seen Dunker’s description or figure, but it seems most
improbable that Thiele’s capensis is the same. Krauss gave the dimensions of
Dunker’s species as 10 X 5:2 lines.
Although there are eight specimens in S. Afr. Museum with their opercula,
no animal has been preserved.
Fusus africanae n. sp.
Figs. 19(k), 20(d)
1925. Thiele. D. Tiefsee Exp., xvii, p. 184, pl. 32 (20), fig. 20 (‘Fusus n. sp.’, sine nom.).
The Valdivia specimen from 35° 16’ S. 22° 26’ E., 155 metres, measured
22°5 < g mm. and consisted of a large protoconch and 3 whorls. The profile
of the whorls is evenly convex, without the bicarinate periphery of juvenile
verruculatus and bonae-sper. Thiele was therefore correct in regarding it as a
distinct species. Unfortunately he did not state the number of ribs and spiral
lirae; judging by his figure there might be 13 or 14 ribs and 4 or 5 lirae on the
end and 3rd whorls.
The Pieter Faure obtained two smaller specimens, 15 X 6:5 mm.; one in
1900 from off west coast of Cape Peninsula, 131-136 fathoms, and another in
1906 from Brown’s Bank (approx. 363° S. 21° E.), 80-100 fathoms (S. Afr.
Mus. nos. A8826 and A8610 Type).
Aperture (incl. canal) a little longer than spire. Protoconch 24 whorls,
diam. 3, alt. 2°75-3 mm., smooth, with 2-3 feeble growth-lines (scarcely ribs)
on outer lip. First postnatal whorl beginning abruptly, and not quite flush
with outer lip of protoconch; 14 low feeble axial ribs on 1st and 15 on 2nd
whorl (slightly irregular in A8826 owing to injury), 3rd whorl incomplete;
crossed by 4 spiral lirae of nearly equal strength (the uppermost one slightly
weaker) on ist and 2nd whorls, a finer one between 1st lira and suture, and
indications of a fine intermediary between each pair of main lirae (more
distinct in A8826 than in A8610), also a lira below the 4th but partly obscured
by suture of following whorl; intersections very slightly nodulose; on base of
last whorl c. 10 main lirae, with finer intermediaries. Columella curved, no
visible pleats, canal slightly curved, a little longer than rest of aperture.
Operculum oval, apex incurved, 3:5 X 2 mm.
6
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 97
The Fisheries Survey vessel Africana obtained (1948) a larger example
Peet, mm., from 34° 35'S. 19 14°. E., 66 metres (AFR. 864 E.).
Protoconch 24 whorls, diam and alt. 3 mm., smooth, with 2-3 feeble lines
of growth on outer lip. Postnatal whorls 5; 1st beginning abruptly and not
quite flush with outer lip of protoconch; axial ribs on 1st whorl obscure,
possibly 14, on 2nd whorl 15 (16) also feeble, on 3rd whorl 16 (17), on 4th
17 (18), thereafter evanescent; crossed by predominant spiral lirae, 5 on Ist
whorl, the lower 3 strongest, a finer one between Ist lira and suture, and below
the 5th another partly obscured by suture of following whorl; on 2nd and
following whorls 6 lirae, the 3 lowest strongest on 2nd and 3rd whorls, the 4
lowest on 4th and 5th whorls, one fine intermediary between each pair of
stronger lirae; on base of last whorl 12 (and some obscure ones on rostrum)
additional lirae with fine intermediaries; lirae on later whorls distinctly
flattened. A small dentiform parietal callus, no visible columella pleats.
Operculum oval, apex incurved, 9 x 6 mm.
Protoconch glistening white, shell greyish-white, operculum horn
coloured.
A very thin periostracum obscured by a thin layer of sponge; and with
numerous oval horny capsules (maj. diam. 0-5 mm.) glued firmly to the shell
(not belonging to this species, or any other Fasciolariid as they are not
stalked).
Radula with c. 190 rows, central plate tricuspid, lateral plate with 11 cusps
(excl. the tiny inner denticle), the innermost 2 strong, the others alternately
smaller and larger, the 10th cusp rather strong and the outermost one the
smallest; the arrangement is not quite symmetrical on the two sides, nor on
successive plates on the same side, particularly so on the left side; the right side
as shown in fig. 19(k) seems to be the most usual arrangement.
Radula of the two juv. P.F. specimens with 105-110 rows, lateral plate
with 5 unequal cusps.
Remarks. The Africana specimen is separately described. to show the slight
difference in detail of the spiral lirae; viewing it side by side with the other two
there is no doubt they are conspecific; and there seems little doubt that they
are the same as the Valdivia example.
The 25 mm. shell in bad condition figured by Thiele (loc. cit., fig. 19)
from the same locality has a smaller protoconch and seems to be referable to
verruculatus.
The four known examples are all from moderate depths on the southern
and south-western slopes of the Agulhas Bank, and it is surprising that no
others have been obtained. From the size of the protoconch one would suspect
an adult at least as large as verruculatus.
The radulae of all three specimens are remarkable for the inequality of
the cusps on the lateral plate, especially noticeable in the large Africana specimen,
thus resembling the radula of Peristernia more than that typical of Fusus.
98 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fusus albinus A. Adams
1855. A. Adams. Proc. Zool. Soc. Lond., p. 222.
1880. Sowerby. Thes. Conch., p. 80, pl. 7 (411), fig. 72.
1903. Von Martens. D. Tiefsee Exp., vii, p. 9, pl. 2, fig. 9 (appressus).
1903. Thiele. ibid., p. 169, pl. 9 (4), fig. 61 (radula) (appressus).
1912. Dautzenberg. Ann. Inst. ocean., V, pt. 3, p. 28.
Adams described ‘a large white solid species with moderately long beak,
and with longitudinal rounded rib-like plicae which are obsolete at the sutures’
from Ichaboe Island, north of Liideritzbucht.
Von Martens compared his species, 101 X 40 mm. from Great Fish Bay
(Angola), 16 fathoms, with albinus, but concluded that the two were distinct.
Dautzenberg recorded albinus from Mossamedes, 15-20 metres, and
Praya Amelia, 15-35 metres, but made no mention of appressus.
? Fasciolaria holcophorus n. sp.
Fig. 21
Fusiform, aperture subequal to spire. Profile of whorls convex, without
shoulder. Protoconch 14 whorls, diam 0-6, alt. 0-5 mm., smooth, with 4 or 5
faint axial ribs on last half whorl, junction with Ist post-
natal whorl indistinct. Postnatal whorls 5; axial ribs 10-11
on ist whorl, 8 on and, 7 on each of the 3rd, 4th and 5th
whorls, from suture to suture and strong on early whorls,
but from later part of 4th whorl becoming feeble and
causing mere undulations on the periphery of 5th whorl,
obsolete on base; crossed by spiral striae, 4 on 1st whorl,
7 on and, 8 on 3rd, 9 on 4th and 10 on 5th, about 16
additional on base, those on rostrum almost parallel with
the canal, all striae regularly spaced except 2 or 3 fine
intermediaries on base. Columella gently curved, with 3
pleats, the lowest one feeble. Canal straight (tip broken).
Periostracum thin, smooth. I1 X 4 mm.
White, periostracum pale buff.
Off Cape St. Blaize, 125 fathoms, 1 dead (S. Afr. Mus.
no. A881Q9. P.F. coll.).
Remarks. In the absence of the radula the generic
position of this pretty little shell is quite uncertain. There
is a somewhat fanciful resemblance to Ptychatractus, which
Fic. 21. . : te
iprtiy stat Thiele (1929) removed from the Fasciolariidae to the
Fasciolaria ? holco- :
phorus n. sp. Vasidae.
Fam. NASSIDAE
Gen. NassA Lam.
1799. Lamarck. Mem. Soc. H. N. Paris, p. 71 (non Nassa Bolten, 1798).
1806, Duméril. ool. Analyt., p. 166 (Nassarius).
a
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 99
1916. Tredale. Proc. Mal. Soc., xii, p. 82 (Nassarius).
1928. Tomlin. Ann. S. Afr. Mus., xxv, p. 313 (Nassarius).
1929. Thiele. Handbuch, 1, p. 322 (incl. Desmoulea [sic]).
1931. Lebour. 7. Mar. Biol. Assoc., xvii, p. 797 (eggs and larva).
1936. Peile. Proc. Mal. Soc., xxii, p. 139 (radula).
1939. id. ibid., xxii, p. 276 (radula).
Remarks. Neither Nassarius Duméril 1806 nor Nassaria Link 1807 (Bucci-
nidae) are to be rejected on account of the similarity of their termination (Rules
Zool. Nomencl. Art. 36. Rec.), though Iredale thought otherwise; but as they
are very liable to confusion, no apology is made for reverting to Nassa Lam;
Thiele accepted it, several years after Iredale’s proposed alteration; and
everyone knows the distinctive facies of a “Nassa’. As regards subgenera, until
some agreement is reached on their definition, they are better ignored, at least
so far as South African species are concerned. These are all typically ‘Nassa’,
with the one exception of kraussiana, the radula of which is also distinctive.
South African ‘beach-conchology’ has run rampant in this genus; and when
once a species has acquired synonyms it seems to attract more synonyms, e.g.
capensis and kochiana (cf. Tomlin). Most of Turton’s ‘n. spp.’ will probably
prove to be synonyms, but without actual examination of his material one
can only suggest possible or likely synonymy. He took no notice of Tomlin’s
1928 paper. :
Nassa analogica Sow.
Fig. 22(a)
1853. A. Adams. Proc. Zool. Soc. Lond. (for 1851), p. 113 (trifasciata, non Gmelin).
1903 (8th July). Sowerby. Mar. Invest. S. Afr., uu, p. 219, pl. 4, fig. 3.
1903. id. ibid., p. 228, pl. 4, fig. 2 (trifasciata Adams).
1903 (18th Dec.). Von Martens. D. Tiefsee Exp., vii, p. 27, pl. 3, fig. 18 (circumtexta).
1903. Thiele. ibid., p. 167, pl. 9 (4), fig. 52 (radula) (circumtexta).
1906. Smith. Ann. Natal Mus., 1, p. 36 (circumtexta and analogica).
1910. Dautzenberg. Act. Soc. Linn. Bordeaux, p. 55 (gallandiana Fischer).
1912. id. Ann. Inst. ocean., vol. V, fasc. 3, p. 33 (trifasciata Adams).
1923. Odhner. Géteb. K. Vet. Handl., xxvi, p. 6 (trifasciata Adams).
1925. Thiele. D. Tiefsee Exp., xvii, p. 182 (circumtexta).
1928. Tomlin. loc. cit., p. 316 (circumtexta).
1932. Turton. Mar. Sh. Pi. Alfred, p. 59 (trifasciata Adams).
Radula. ‘Thiele gave the number of cusps on the central plate of a radula
4°75 mm. long as 8. In S. Afr. Mus. specimens of radulae 3:5-4:5 mm. long,
there are 65-75 rows, central plate with 11-13 (rarely 10) cusps, slightly
variable in size znier se, with a minute one externally on both sides, no inter-
mediate plate, outer cusp of the lateral plate with a slight bulge (scarcely a
denticle) on inner margin, but sometimes obscure, inner cusp rather slender.
No difference in the radulae of the ‘trifasciata’ and analogica forms.
The 1st whorl always, and usually also the 2nd whorl, appear to have
only spiral lirae; but axial ribs may be developed on the 3rd and 4th whorls,
or on the 4th and 5th, or on the 5th and 6th, or on all these whorls (3-6). The
number of ribs is 14-15 on 3rd whorl, c. 20 on 5th, and 22-24 on 6th.
oO ANNALS OF THE SOUTH AFRICAN MUSEUM
Sl A) el
WH) wi
A
—
Ae.
SI<p) Siew
Saas 7) Sl
Central and lateral radula plates of Nassa (a) analogica Sow.; (b) kochiana (Dnkr.); (c) speciosa
A. Adams; (d) desmoulioides Sow.; (e) babylonica Watson; (jf) eusulcata Sow.; (g) gemmulata
(Lam.); (h) bicallosa Smith; (i) arcularia Linn.; (j) coronata Brug.; (k) kraussiana (Dnkr.);
(Ll) Demoulia abbreviata (Gmelin).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA IOI
The ribs are distinctly oblique (retractive), and when present on the 6th
whorl they cross the base as far as the columella glaze.
Periostracum thin, minutely fimbriate in some well-preserved specimens,
pale grey or buff.
Operculum triangularly oval, margins entire.
Dead: west coast Cape Peninsula, Port Alfred; also Great Fish Bay
(Angola) (Thiele).
Living (and dead): ‘trifasciata’ form: from off East London to False Bay
and off Cape Point, 26-58 fathoms (von Martens, Sowerby, and 8S. Afr. Mus.
P.F. coll.) ; ‘analogica’ form: from False Bay around Cape Point to the Saldanha
Bay area, 22-80 fathoms (S. Afr. Mus. P.F. coll.); St. Helena Bay, 27 metres
Seeee)-@. 1). 26" 39'S 15° E., 55 metres (s.s. Africana, per U.C.T.).
Remarks. ‘The extreme forms: a fully plicate ‘trifasciata’ and a spirally
lirate analogica, are very different in appearance. Sowerby said (1903, p. 220)
the two forms ‘when separated’ showed very little variation; the difficulty is to
separate them!
In analogica the spiral lirae are always flat-topped, but their width varies
according as they are separated by narrow striae or flat-bottomed grooves,
which latter may sometimes be almost as wide as the lirae. The width of the
spiral lirae appears to be less variable in the plicate form.
The Pieter Faure material in the S. Afr. Museum shows that the more or
less plicate form is found on the south coast as far west as False Bay and Cape
Point, but not on the west coast (a beach-worn specimen from Table Bay is an
exception) ; the lirate form is more characteristic of the west coast, but occurs
throughout the south coast area.
Both von Martens and Smith regard Adams’s locality (Vigo Bay, Spain)
as erroneous.
Dautzenberg (1912) regards gallandiana Fischer 1862 as synonymous with
triafasciata Adams, with distribution: mouth of the Congo River, 25 metres,
and (loc. cit. 1910) Lagos and Senegal. The identity of the Angolan, and
especially the west African examples, with South African examples requires
investigation.
Nassa pyramidalis (A. Adams)
1853. A. Adams. Proc. Zool. Soc. Lond. (for 1851), p. 113 (Desmoulea p.).
1886. Sowerby. 7. Conch., v, p. 4 (Desmoulea p.).
1900. id. Proc. Mal. Soc., iv, p. 2, pl. 1, fig. 5 (filmerae).
1928. Tomlin. loc. cit., p. 318 (j/ilmerae) and p. 325 (pyramidalis).
1932. Turton. Mar. Sh. Pt. Alfred, p. 59, pl. 14, nos. 435, 436, 437 (pyramidalis, and vars.
affiinis and punctilineata) ; and p. 59 ( filmerae).
1932. id. 7. Conch., xix, p. 370 (rufanensis nom. nov. for affinis preocc. Sow. 1832).
Pyramidal. Protoconch 3 whorls, diam. and alt. 0-75-0-8 mm., smooth,
glossy. Postnatal whorls 7; axial ribs 13-14 on Ist, increasing to 14-15, from
suture to suture, straight or slightly oblique (protractive), more strongly marked
on early whorls, and usually evanescent on last half of 7th whorl; crossed by
102 ANNALS OF THE SOUTH AFRICAN MUSEUM
spiral lirae, 4 on 1st whorl, 5 on 2nd, 6 on 3rd and increasing to 10-11 on last
whorl, 12-14 additional lirae on base and 7—8 on rostrum. On later whorls the
lirae become flattened and broader than the intervening striae, and the surface
of the whorls is better described as being smooth with impressed spiral striae; a
very characteristic sculpture. Internal parietal callus nodular, columella
smooth, anteriorly subcarinate, columella glaze not extensive, thin. Outer lip
internally plicate, and end plica posteriorly and anteriorly dentiform. 22
(protoconch and first 2 whorls missing) * 13 mm.
Operculum and radula?
Cream with orange-brown irregular marks, more or less connected to
form a sutural band, or with a series of dots along some of the lirae, aperture
and columella more or less brownish, apex in worn specimens sometimes
violaceous.
Port Elizabeth, Port Alfred, Pondoland. Mouth of Gouritz River, and
Jeffreys Bay (S. Afr. Mus.). Algoa Bay, 10 fathoms, off Cape St. Blaize, 28
fathoms, off Cove Rock (East London area), 80-130 fathoms, off Umhloti
River (Natal), 40 fathoms (S. Afr. Mus. PF. coll.). 30° 47 S5}3@ seg ee
24 fathoms (U.C.T.).
Remarks. Only dead specimens known, though one of the Cape St. Blaize
specimens, and the 6 juveniles from Umhloti, obtained by the Pieter Faure, were
quite fresh. U.C.T. obtained one dead shell in Algoa Bay, and one dead but
fresh specimen off Scottburgh, Natal.
There are 3 cotypes of filmerae from Pondoland (don. Dr. H. Becker) in
S. Afr. Museum; it seems strange that Tomlin did not definitely unite this
with pyramidalis.
Nassa babylonica Watson
Figs. 22(e), 23(e)
1882. Watson. 7. Linn. Soc. Lond., xvi, p. 366.
1886. id. Challenger Rep., xv, p. 185, pl. xi, fig. 8 (with protoconch and operculum).
1899. Smith. Ann. Mag. Nat. Hist. (7), iv, p. 243 (diluta).
1901. Illustr. Zool. Investigator. Moll., pl. xi, figs. 3, 3a (diluta).
1901. Melvill & Standen. Proc. Zool. Soc. Lond., ii, p. 409.
1903. Von Martens. D. Tiefsee Exp., vii, p. 100, pl. 3, figs. 7, 8.
1903. Thiele. ibid., p. 167, pl. 9 (4), fig. 53 (radula).
1925. id. ibid., xvii, p. 183.
1928. Tomlin. loc, cit., p. 314.
Turreted, with square shoulders. Protoconch 34-34 whorls, diam. and
alt. 1 mm., smooth, the 3rd whorl carinate below middle of whorl, the carina
descending into the suture on last quarter whorl. Postnatal whorls 5; axial
ribs 11-12 on Ist whorl, increasing to 14-15 on last whorl, sharply tubercular
at top, slightly indented immediately below; spiral lirae feeble, 3-4 traceable
on lower part of 4th and 5th whorl, with 3-4 stronger additional ones on base.
Outer lip feebly denticulate within. 0-45 x 0:23 in. (Watson); 10 X 5 mm.
(S. Afr. Mus.); 12 x 7 mm. (Smith: diluta).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 103
Operculum triangularly oval (more triangular than in Watson’s figure),
not quite twice as long as broad, margins entire.
Radula with 60-65 rows, central plate strongly concave in front, with 8-9
(10) cusps, slightly varying in size inter se, the middle one usually the largest,
the outermost one or two on either side minute, lateral plate with rather short
cusps, the inner one stout, no intermediate plate.
Off Cape Natal (Durban), 400 fathoms, 24 specimens, most of them dead
feseuir. Mus. P.F. coll.).
Distribution. East Africa. 1,134-1,644 metres (von Martens); Gulf of
Oman and Karachi, 37-80 fathoms (Melvill and Standen); off Ceylon, 597
and 753 fathoms (Smith); Philippine Islands, 375 fathoms (Watson).
Remarks. Von Martens illustrated plump and slender forms. In the
present lot there are two broken specimens with 4 whorls measuring
oe 3, 0 7) ID.
Smith’s diluta has one or two fewer ribs, but can scarcely be regarded as
distinct.
Only one specimen was available for Tomlin’s inspection; the other
examples have since been found in a bottom sample from the same locality.
For comparison of protoconch with that of bicallosa, see latter p. 108.
Nassa capensis (Dnkr.)
Fig. 23(a)
1880. Von Martens. Mauritius G Seychellen, p. 243.
1928. Tomlin. loc. cit., p. 315 (references and synonymy).
1932. Turton. Mar. Sh. Pt. Alfred, p. 55, pl. 12, no. 412 (kraussi).
1932. id. ibid., p. 58, pl. 13, no. 425 (ordinaria).
Protoconch 24 whorls, diam 0-6, alt. 0-75 mm., smooth. Postnatal whorls
7; axial ribs (9g) 10 on ist whorl, 10-11 on 2nd, 11-12 on 3rd, 12-13 (14) on
last whorl, suture to suture, oblique, protractive, evanescent on base; crossed
by spiral lirae 4 (5) on 2nd—4th whorls, obscure on ist and usually not con-
tinued on to 5th, always obsolete on 6th and 7th leaving the ribs and intervals
perfectly smooth; 2 rather broad flat spiral lirae on base anteriorly and 6
striae on rostrum. Internal parietal callus cariniform, columella smooth,
carinate at anterior end, columella glaze not extensive, thin. Outer lip with
varix in adult, internally plicate (but often feebly). 16-5 x 6 mm.
Operculum and radula?
Cream or buff, speckled or dappled, with a more or less marked brown
band on lower part of base, bordered above by a disconnected series of dashes
(one in each interval between the ribs) which appears just above suture in
preceding whorls; in juveniles protoconch and rst whorl brown, following
whorls glistening white; sometimes pure white (but ? faded), or unicolorous
yellowish, amber, ochraceous or brown (serotina); anterior canal often rusty
brown.
104 ANNALS OF THE SOUTH AFRICAN MUSEUM
False Bay to East London and Natal (Tomlin, S. Afr. Mus.); also (small
form) 'Tongaat (Natal).
Distribution. Mauritius and Réunion (von Martens).
Remarks. Plump and slender forms occur, e.g.: 10 X 5 mm. and
10 X 4mm. The ribs on successive whorls are more or less in an axial line,
sometimes very distinctly so (resembling a Scalaria).
Protoconchs and juveniles were collected at Still Bay by the late Dr. Muir.
It is remarkable that no living specimens have been collected.
The Mauritius and Réunion records should be checked, also Bisacchi’s
(1930) record of pulchella from Suez.
Fic. 23.
Protoconchs of Nassa (a) capensis (Dnkr.); (6) kochiana (Dnkr.) ; (c) desmoulioides Sow.; (d) bicallosa
Smith; (e) babylonica Watson; (f ) kraussiana (Dnkr.).
Nassa kochiana (Dnkr.)
Figs. 22(b), 23(b)
1903. Von Martens. D. Tiefsee Exp., vii, p. 28 (crawford).
1925. Thiele. ibid., xvii, p. 182 (limata Chemn.).
1928. ‘Tomlin. loc. cit., p. 319 (references and synonymy) and p. 327.
1932. Turton. Mar. Sh. Pt. Alfred, p. 56, pl. 13, no. 417 (eucosmia), and p. 57, pl. 13, no. 419
(carinata).
Protoconch 24 whorls, diam. 0-6, alt. 0:75 mm., smooth. Postnatal
whorls, 7; axial ribs 12-13 on 1st whorl, 13-14 on 2nd, 14-15 on 3rd, increasing
to 15 or 16 to 18 on last whorl, from suture to suture, slightly curved and
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 105
protractive, continued across base; crossed by spiral lirae 4 on 1st whorl, 5 (6)
on 2nd, 6 (7) on 3rd, increasing to 13-14 on last whorl, 2 or 3 of the lirae on
early whorls and 4 or 5 on the later whorls near the suture are fine and narrower
than the others; 5 (6) additional lirae on base, sometimes with finer inter-
-mediaries, 6 striae on rostrum. Internal parietal callus cariniform, columella
smooth but sometimes feebly granulate, carinate at anterior end, columellar
glaze not extensive, thin. Outer lip with varix in adult, internally plicate.
14 X 6mm.
Operculum oval, margins entire.
Cream or buff with faint dappling or spiral lines, chiefly on base, with a
series of brown spots forming a broken narrow spiral band just below periphery
from upper end of aperture on last whorl, on earlier whorls just above suture,
sometimes with a white band above the dark band (spurca and poecilostoma) ; or
unicolorous yellowish, ochraceous, brown, or pinkish (coccinea); in juveniles
protoconch and early whorls glassy white with indications (on 2nd and 3rd
whorl) of dappling and the brown spiral line; anterior canal often rusty brown;
or with 9-10 pale brown narrow spiral bands on last whorl (incl. base), the
ends of which appear on the outer lip varix as double lines (crawford).
Radula with c. 60 rows, central plate strongly concave in front, with 9
cusps and a minute one externally on either side, inner cusp of lateral plate
moderately stout, no intermediate plate.
Fossil: raised beach Algoa Bay.
False Bay to Port Alfred. The only record from the west coast (Tryon:
Table Bay) is scarcely acceptable.
Living: False Bay, 24 metres (U.C.T.).
Remarks. ‘Thiele (1925) disagreed with von Martens’s (1903) identification
of shells from St. Francis Bay as crawfordi, preferring to identify them with
limata Chemn. (Mediterranean, Madeira, Canaries, Gape Verde Is., and West
Africa); von Martens is more likely to have been correct.
Dautzenberg (1912, Ann. Inst. ocean., vol. 5, fasc. 3, p. 31) recorded poecilo-
stoma Smith from Mossamedes, littoral and 15-20 metres. I think these
specimens should be re-examined.
The obliquity of the ribs varies, and may vary from whorl to whorl on the
same shell. Also there are plump and slender forms.
Comparison of kochiana with European incrassata at once shows the
differences: in the latter the axial ribs are retractive, the number of spiral lirae
on the 6th whorl is only 6, the intersections are more nodulose, the profile is
undulate (notched in kochiana), and there are no fine lirae below the suture,
or at most only one and that one is only very slightly narrower than the
following lirae.
Nassa muiri n. sp.
1932. Turton. Mar. Sh. Pt. Alfred, p. 58, pl. 13, no. 428 (microstoma, non Pease) and no. 429
(ambigua, non Mont.).
106 ANNALS OF THE SOUTH AFRICAN MUSEUM
Protoconch 24 whorls, diam. 1-1-25, alt. 0-75-o-8 mm., smooth. Post-
natal whorls 4, profile rather strongly convex; axial ribs 15 on 1st whorl, 18-19
on 2nd, 21-23 on 3rd and 4th, but often obscure on 4th and only c. 17-18
countable, sometimes only 17 on 2nd and 3rd, and 18 on 4th whorl, straight
(or nearly so) and retractive; crossed by spiral lirae 6 on 1st whorl, 7 on 2nd,
8 on grd, g-10 on 4th, intersections here and there irregularly nodulose, 7-8
additional lirae on base, 4—5 striae on rostrum. Columella smooth, angulate
anteriorly, columellar glaze not extensive; parietal callus feeble; outer tip with
varix, internally more or less plicate. 10 X 6-5 mm. (Turton gave ‘c. 16 mm.’
for microstoma, but the line alongside the figure is only 11 mm.)
Operculum oval, margins entire.
Pale buff, with irregular orange-brown marks giving an impression of
more or less distinct flames, chiefly around upper half of whorl, which may
thus be nearly uniformly brown; two or three marks on outer lip varix;
anterior canal orange-brown.
Radula with 60-65 rows, central plate strongly concave in front, 8-g (10)
cusps, with a small or minute one externally on either side, inner cusp of
lateral plate moderately stout, no intermediate plate.
Port Alfred (Turton, also S$. Afr. Mus. one example from Turton labelled
‘quantula’); a series from possibly the same locality (S. Afr. Mus.); Still Bay,
juvenile (protoconch plus 2 whorls), very fresh (S. Afr. Mus. Muir coll.) ; from
stomach of seal caught in False Bay, seven examples (S. Afr. Mus.); Algoa Bay,
52 fathoms, and off East London, 32 fathoms (S. Afr. Mus. P.F. coll.); False
Bay (34° 18’ S. 18° 29’ E.), 51 metres (U.C.T.); 34° 15'S. 25° 5 ito easy
and 33°S. 26° 11’ E. (off East London), 31 fathoms (U.C.T.).
Remarks. ‘Turton’s photographic figures of what he recorded as possibly
microstoma and ambigua, certainly represent this species. Mr. Salisbury informs
me that it is not the West Indian ambigua.
The straight and slightly retractive axial ribs distinguish it from kochiana.
Named after the late Dr. John Muir, of Riversdale.
I am diffident about introducing another species so close to plebeja Thiele,
but the shells can be distinguished at a glance by the more numerous spiral
lirae in muri. ‘The sulci are consequently narrower than the lirae, which pro-
duces a lirate sculpture; whereas in plebeja the lirae and sulci are subequal,
producing a more cancellate sculpture.
Nassa plebeja Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 182, pl. 32 (20), fig. 9.
Protoconch 24 whorls, diam. 0-75, alt. 0-6 mm., smooth. Postnatal
whorls 44; axial ribs 14 on 1st whorl, increasing to 15 (16 incl. varix) on last
whorl, straight or slightly curved, slightly retractive, obsolete on base of body
whorl; crossed by spiral lirae 4. on 1st-3rd whorls, 5 on 4th, on 3rd and 4th
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 107
whorls an additional finer lira appears between the suture and the uppermost
lira, 4 additional rather sharp lirae on base and 5-6 on rostrum; intersections
with ribs slightly nodulose. Columella angulate anteriorly, columellar glaze
narrow; parietal callus feeble (distinct in Thiele’s figure) ; outer lip with varix,
plicate within. 7 X 4 mm. Thiele: 6-25 X 3-5 mm.
Uniform greyish. Brown above and below the periphery, anterior canal
darker (Thiele).
St. Francis Bay, 80 metres, and Algoa Bay (Thiele). Algoa Bay 51 fathoms,
two dead (S. Afr. Mus. P.F. coll.). False Bay (U.C.T.).
Distribution. Great Fish Bay, Angola (Thiele).
Remarks. The single U.C.T. specimen and the two P.F. specimens appear
to agree with plebeja, assuming that Thiele’s figure is correct; his description
does not give the number of axial ribs or spiral lirae.
The identity of the Angolan specimens is provisionally accepted.
cf. Odhner’s record of ambigua Mont. from Port Alexander (1923. Goteb. K.
Vet. Handl., xxvi, 7, p. 14).
Nassa bicallosa Smith
Figs. 22(h), 23(d)
1876. Smith. 7. Linn. Soc. Lond., xii, p. 543, pl. 30, fig. 1.
1928. Tomlin. loc. cit., p. 313 (algida, non Rve., part: S. Afr. Mus. no. A6398).
1928. id. ibid., p. 314.
Protoconch 34 whorls, diam. 1, alt. 0-75-o-8 mm., smooth, a faint peri-
pheral keel on last whorl and half. Postnatal whorls 6 (7 in two specimens) ;
profile of whorls straight or nearly so, squarely shouldered at top; axial ribs
13-14 on early whorls, increasing to 15-16 or 17 on later whorls, where some
of them are often more or less duplicated, from suture to suture and distinctly
oblique (retractive) from ist to 5th whorls, on later part of 5th and on 6th
becoming evanescent except as knobs at top of whorl, sometimes strongly,
sometimes feebly developed; one spiral stria marking off the coronal knobs on
all whorls, but evanescent on last whorl, 6—7 striae on lower part of base and
3-4 on rostrum. External parietal callus opposing tooth on outer lip nodular
(in adult), labral sinus deep, internal callus dentiform or subcarinate; colu-
mella anteriorly carinate and denticulate, glaze not extending over base, its
edge free. Outer lip internally plicate. 26 x 15 mm. (6 whorls), 27 X 15 mm.
(7 whorls).
Operculum triangularly ovate, serrate on both margins (when not
corroded), 7-8 X 5 mm.
Living examples from 24 fathoms varying from buff, slightly stained with
orange, to reddish-brown, partly coated with some black substance, aperture
brownish within, outer lip white, operculum amber or dark brown.
Specimens from 14 fathoms varying from reddish or chestnut brown to
almost black, most specimens more or less corroded and covered with some
108 ANNALS OF THE SOUTH AFRICAN MUSEUM
(? algal) substance, chiefly on the upper side while the lower side remains
polished.
Radula with c. 75 rows, central plate with 16 cusps, outermost one or two
on either side minute, intermediate plate oval, lateral plate without denticles
between the 2 cusps.
Cape Natal (Durban) (Smith). Off Tongaat, 36 fathoms, one juv. dead,
and off Umhlanga, 22-26 fathoms one juv. dead; off Umkomaas, 40 fathoms,
3 protoconchs (S. Afr. Mus. P.F. coll.).
Living: off Tugela River, 12-14 and 24 fathoms (Tomlin, S. Afr. Mus.
Pal coll®):
Remarks. ‘Tomlin, by a slip, referred some of the specimens (S. Afr. Mus.
no. A6398) dredged in 12-14 fathoms to “‘algida’; they are clearly the same as
those dredged in 24 fathoms (A6399).
The specimens from the deeper water are not corroded and are quite
clean except for traces of foreign matter on the upper whorls; they are also
more strongly coronate than most of the shallower water examples. The
operculum of the latter is also more or less corroded to an irregular oval shape.
The double columella callus may serve to distinguish this species from
glans-suturalis, but is found in other species e.g. coronata.
The three very juvenile examples, consisting of protoconch and first two
postnatal whorls, one of them quite fresh and translucent, show that the species
is living as far south as Umkomaas.
Apart from the 1st and 2nd postnatal whorls, the protoconch would be
indistinguishable from that of babylonica; the straight-sided early whorls are
very similar in the two species, but bzcallosa has one or two more ribs, and the
tops of the ribs are not so mucronate as in babylonica. The profile of the later
whorls becomes gradually oblique in bicallosa, producing a pyramidal shell,
whereas in babylonica it remains essentially vertical, and the adult shell is
turreted.
Nassa glans (Linn.)
1758. Linne. Syst. Nat., 1oth ed., p. 737, sp. 394 (Buccinum g.).
1822. Lamarck. Anim. sans. Vert., vii, p. 269 (Buccinum suturale).
1859. Chenu. Man. Conch., 1, fig. 771.
1880. Von Martens. Mauritius G Seychellen, p. 242 (suturalis).
1886. Watson. Challenger Rep., xv, p. 179 (references).
? igor. Smith. 7. Conch., x, p. 111, pl. 1, fig. 17 (algida, non Rve.).
1928. Tomlin. loc. cit., p. 313 (algida, non Rve., part: S. Afr. Mus. no. 14039).
1928. id. ibid., p. 325 (suturalis).
1952. Satyamurti. Bull. Madras Govt. Mus., n.s. I, ii, 6, p. 184, pl. 17, figs. 7 a, 6 (suturalis).
1956. Day & Morgans. Ann. Natal Mus., xiii, p. 306 (listed, as algidus, non Rve.).
Protoconch 14 (2) whorls, smooth. Postnatal whorls 6; axial ribs c. 18-22
on early whorls, from suture to suture, but evanescent on 6th whorl except as
a series of coronal nodules; crossed by 3 spiral striae dividing the ribs into 4
approximately equal-sized nodules, evanescent as impressed striae on 5th
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 109
whorl, but often indicated by thin coloured lines, a 4th stria partly visible in the
suture; 4 additional lines on base, the lower 2 of which are again impressed
striae, 6-8 striae on rostrum. Internal parietal callus strongly cariniform,
columella carinate at anterior end, glaze not extensive, edge free on rostrum.
Outer lip thin, with strong tooth posteriorly opposite the parietal callus,
forming a deep sinus, internally non-plicate. 25 x 13 mm. (S. Afr. Mus.);
31 X 18 mm. (Smith).
Operculum triangularly ovate, serrate on both margins, 3 X 2°5 mm. in
aperture 10 mm. in shell 20 mm. long.
Yellowish with diffuse orange blotches or flames, early whorls pinkish,
protoconch crimson, the spiral striae and lines orange-brown (dead specimens).
Animal pale, spotted with black on siphon and proboscis, tentacles pale.
Radula with c. 70 rows, central plate with 10 cusps, intermediate plate
oval, lateral plate without denticles between the cusps.
Durban (Smith, ‘Tomlin, S. Afr. Mus.).
Distribution. Zanzibar (S. Afr. Mus.), Mauritius, Ile de France, Indo-
Pacific.
Remarks. N. suturalis is regarded as a small variety of glans (Watson,
locs.cit.).
One of the shells described above was identified many years ago by
J. H. Ponsonby, who doubted its South African provenance; later Tomlin
confirmed the identity of this shell. Tomlin also identified 4 shells from Durban
(S. Afr. Mus. no. 14039) as algida Rve. I cannot agree because the sculpture
of the early whorls is the same as in the shell labelled suturalis, and the coloration
is similar except that the orange-brown spiral lines are faded or worn away. I
suggest also that Smith’s identification was erroneous, and that the Australian
algida be deleted from the South African fauna-list.
S. Afr. Museum has 2 examples of glans from the Philippines (ex Ross-
Frames coll.): 7 whorls 44 x 23 mm., and 6 whorls 39 x 21 mm. Also 2 from
Zanzibar (E. L. Layard, H.M.S. Castor, coll. 1856); 6 whorls 30 x 16 mm.
and 5 whorls 21 x 11 mm. In these the 4th spiral line is visible throughout,
nowhere covered by the suture; otherwise there is no difference, except the
large examples are feebly plicate within the aperture.
Nassa coronata Brug.
Fig. 22(7)
1789. Bruguiére. Encycl. Meth. Vers, 1, p. 276 (Buccinum c.).
1880. Von Martens. Mauritius G Seychellen, p. 242.
1928. Tomlin. loc. cit., pp. 317 and 327.
Protoconch 14 whorls, smooth. Postnatal whorls 6; axial ribs 20-21 on
ist whorl, 22-23 on 2nd, 23-24 on 3rd, then decreasing to 18 on 4th, 10-12
on 5th and 6th whorls, suture to suture and straight or very slightly oblique
(retractive) on Ist to 4th whorls, on 5th and 6th evanescent except as rounded
110 ANNALS OF THE SOUTH AFRICAN MUSEUM
nodules forming a coronal shoulder; crossed by 3 spiral striae, one above and
2 below, the and and 3rd closer together than the 1st and and, demarcating
3 series of nodules, rounded in the uppermost series, axially elongate oblongs
in the middle series and squarish areas in the third series; spirals continued on
to 5th whorl but thereafter obsolete; 3 striae on lower part of base, 3-4 on
rostrum. External parietal callus nodular, internal callus dentiform or sub-
carinate, labral sinus deep; columella subcarinate anteriorly and with 2-3
feeble plicae, glaze extensive, thickened in adult. Outer lip internally plicate.
32 X 20 mm.
Operculum subtriangular, 5-8 serrations on both margins, 6 x 6 mm. in
28 mm. shell.
Grey or bluish-grey, with cream coloured coronal knobs and a spiral band
in middle of whorl, a narrower band lower down on base; outer lip externally
and internally, and columella glaze white, aperture within brownish with the
external pale bands showing through.
Radula with 65-75 rows, central plate with 10-12 cusps, intermediate
plate piriform, lateral plate without denticles between the 2 cusps.
Fossil: raised beach, 375 ft. alt., Durban-Umgeni (Geol. Surv.).
Natal (Krauss, Tomlin, 8. Afr. Mus.).
Living: Delagoa Bay (U.W.); Inhambane (U.C.T.).
Distribution. Mauritius, Madagascar, Aden, Indo-Pacific.
Nassa marganitifer (Dnkr.)
1928. Tomlin. loc. cit., p. 321.
Protoconch 14 (2) whorls, smooth. Postnatal whorls 7; axial aribs 17-18
on 1st whorl, increasing to 24-26 on last, from suture to suture, straight,
slightly oblique (retractive); crossed by one open groove on Ist whorl, 2 on
end and 3rd, and 3 on 4th—6th whorls, with a feebler 4th groove on later part
of 6th and on 7th whorl, dividing the ribs into 3 series of rounded nodules, 4
series on 6th and 7th whorls; the uppermost groove below the subsutural
nodules conspicuously wider than the other grooves; ribs on base extending to
columella glaze and divided by 7 grooves into nodules. External parietal
callus bulbous but not prominent, internal callus strongly cariniform, columella
nodulose, carinate anteriorly, glaze not extensive but rather thick, edge on
rostrum free. Outer lip feebly toothed posteriorly, internally plicate, externally
thickened, more or less varicoid. 27 X 14 mm.
Operculum triangularly oval, margins undulate (probably serrate when
fresh), 7 X 4 mm. in 26 mm. shell.
White with greyish shading and slaty-grey or purplish-brown irregular
blotches and markings, a more or less continuous band of same colour around
middle of last whorl (concealed by suture in earlier whorls), and another less
well-marked below suture, columella and outer lip white, aperture within with
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA PED
i-3 (usually 3) purplish bands, the one corresponding with the external
peripheral band always present.
Dead: Durban (Smith); Bartholomew Diaz, Bazarute Islands, Portuguese
East Africa (S. Afr. Mus. coll. Ross-Frames).
Distribution. Red Sea, Ceylon, Indo-Pacific.
Remarks. ‘The Ross-Frames specimens were probably taken alive, though
none of them now possess opercula.
A worn specimen in 8. Afr. Mus. (no. 2623) registered as from Durban,
was identified by Tomlin as the European reticulata Linn. ‘doubtless from
ballast’. It has a sutural and a peripheral dark band, and is so similar to
margaritifer that the following differences between the two species may be given
in case more ‘ballast’ specimens come to hand.
In fresh retzculatus (British specimens) there are only 12 axial ribs on Ist
whorl, increasing to 20 on 7th; on last whorl there are 5 spiral sulci dividing
each rib into 5 nodules, 7 additional sulci on base; the uppermost sulcus is an
open groove (as in margaritifer) but the 1st nodule below the suture is double;
a small nodule separated by a narrow impressed line from the larger (main)
nodule; all the nodules are flat above, sloping below, so that the profile of the
whorl is stepped or feebly serrate (in margaritifer it is only undulate). No
external parietal callus and internal callus very feeble, columella nodulose
only at anterior end, glaze more extensive. All these features are traceable in
the ‘ballast’ specimen and confirm Tomlin’s identification.
Braga (1952) has recorded albescens Dnkr. from Mozambique, and Thiele
(1925) described incognita from Dar-es-Salaam, both of which might usefully
be compared with the present species.
1848.
1877.
1928.
1931.
Nassa fenestrata Marrat
Krauss. Stidafr. Moll., p. 122 (Buccinum marginulatum, non Lam.).
Marrat. New Forms of Nassa, p. 10.
Tomlin. loc. cit., p. 317, also p. 321 (marginulatus).
id. 7. Conch., xix, p. 107.
Protoconch 14 (2) whorls, smooth. Postnatal whorls 5; axial ribs 24 on
Ist whorl, increasing to 26 on 5th, from suture to suture, straight but slightly
oblique (retractive) on later whorls; crossed by 3 spiral striae on 3rd whorl,
dividing the ribs into 4 nodules, on 4th whorl upper (sutural) nodules divided
by an additional stria, and 2 striae between Ist (sutural) and 2nd series of
nodules, producing a narrow lira between the two series; on 5th whorl a and
stria dividing upper nodules into 3, and the narrow lira is repeated between
each pair of nodules, thus broad and narrow nodules alternating; on base 7
eadditional series of nodules (or broad lirae) alternating with narrow lirae.
Internal parietal callus cariniform, columella nodulose anteriorly, glaze strong,
thick and somewhat bulbous. Outer lip reflexed, thickened, glazed, internally
plicate. 20 X 11°5 mm., and plump form 18 X 12 mm,
I12 ANNALS OF THE SOUTH AFRICAN MUSEUM
Operculum and radula?
Cream or buff or greyish, 3 indistinct darker greyish bands: one sub-
sutural, one peripheral, and one at bottom of base; columella glaze and outer
lip white, aperture internally with 3 more or less distinct brownish bands.
Dead: Mozambique (Marrat); Durban (Krauss, Sowerby, 'Tomlin, S. Afr.
Mus.).
Living: Inhambane (S. Afr. Mus.); Delagoa Bay (U.W.).
Distribution. Mauritius (S. Afr. Mus.) ; Red Sea; Philippines and Australia
(Marrat).
Remarks. The Inhambane specimens were evidently taken alive because
the animals have been eaten out by Anthrenus (larval skins remaining in
aperture), but the opercula have been lost.
N. martensi ‘Thiele 1925 from Dar-es-Salaam, should be compared with
this species.
Nassa eusulcata Sow.
Fig. 22( f)
1902. Sowerby. Mar. Invest. S. Afr., ii, p. 94, pl. 2, fig. 8.
1928. Tomlin. loc. cit., p. 317.
Protoconch 2 (24) whorls, smooth. Postnatal whorls 7; axial ribs 13-14
on 1st whorl, increasing to 18 on last, but often on back of outer lip additional
ribs crowded together to a total of 20-26, from suture to suture, slightly oblique
(retractive); crossed by 3 spiral striae on 1st—4th whorl, 4 on 5th, 5 on 6th,
and 6 on 7th whorl, the uppermost stria however from 4th whorl onwards
becoming a well-marked sulcus separating the tops of the ribs as a subsutural
series of nodules; on base 7 (8) striae become stronger and more open sulci
anteriorly, with rather sharp intervening lirae, the intersections slightly
nodular. Internal parietal callus cariniform, followed by columella nodules,
glaze narrow, not spreading over base. Outer lip plicate within. 19 X 10 mm.
Operculum broadly oval, 3 strong serrations on outer, 2 on inner margin,
Ax 8 fom.in 17mm. shell:
Radula with 60-65 rows, central plate with 10 cusps (12 in one of three
specimens), outermost on either side minute, no intermediate plate, lateral
plate without denticles between the 2 cusps.
Living (and dead): off Tugela River, and off O’Neil Peak (Zululand),
40-55 fathoms (Sowerby, Tomlin, S. Afr. Mus. P.F. coll.).
Nassa natalensis Smith
1903. Smith. Proc. Mal. Soc., v, p. 373, pl. 15, fig. 6.
1928. Tomlin. loc. cit., p. 322.
1936. Peile. Proc. Mal. Soc., xxii, p. 140 (radula).
Protoconch ? Postnatal whorls 6, profile angular in middle of whorl;
axial ribs 12 on 2nd whorl, 12-13 on later whorls, strong on early whorls, less
7
id
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 113
so on last whorl, nodular at top below suture and at periphery, continued over
base where they are also nodulous; 6 lirae on base, obscure above, stronger
below. External parietal callus strong, bulbous, internal callus cariniform,
columella smooth, carinate anteriorly, glaze rather broad but not extending
over base, edge free on rostrum. Outer lip thickened, with varix, internally
plicate. 20 X 12-5 mm. (Smith’s figure: 21 mm.)
Operculum ?
Cream or greyish, on body whorl a series of horizontally oblong dark
brown streaks between the ribs, forming a narrow broken band from posterior
end of aperture, mostly concealed by suture in earlier whorls (Smith’s figure
shows the dark band crossing the ribs as well as the intervals) ; Smith mentioned
one specimen as being ‘a rich brown colour with a white line above the middle
of the body whorl’.
Radula (Peile) lateral plate with denticles between the 2 cusps, inner edge
of the inner cusp sometimes serrulate, ? intermediate plate (not mentioned).
Natal (Reeve, Smith, S. Afr. Mus.); Mozambique Island (U.W.).
Nassa arcularia Linn.
Fig. 22(2)
1852. A. Adams. Proc. ool. Soc. Lond. (for 1851), p. 98 (sulcifera = monstrosity).
1880. Von Martens. Mauritius G Seychellen, p. 242 (incl. rumphit Desh. and pullus Lam.).
1928. Tomlin. loc. cit., p. 314, p. 324 (pullus), and p. 327.
1930. Bisacchi. Ann. Mus. Civ. Genoa, lv, p. 44, and p. 47, figs. 1-3 (pullus).
1933. Krige. Tr. Geol. Soc. S. Afr., xxxv (1932), p. 52.
1938. Adams & Leloup. Mem. Mus. Roy. H. N. Belg., Hors Série II, 19, p. 183, pl. 8,
fig. 7a; b\( juv.).
1942. Gravely. Bull. Madras Govt. Mus., n.s. V, no. 2, p. 60 (in key), fig. 11 e. (pulla).
1952. Braga. Anais Est. Zool. Ultramar, vii, 3, p. 74, pl. 3, fig. 2.
Protoconch 14 whorls, smooth. Postnatal whorls 5; axial ribs 19-20 on
ist and 2nd whorls, 16-17 on grd, 15 on 4th, 13-15 on 5th, suture to suture,
slightly oblique (retractive), continued on base except on last portion of 5th
whorl where there are 4-6 oblique costae followed by the labral varix; crossed
by 3 spiral striae, equidistant, the uppermost a little distance below the coronal
knobs, 5-6 striae on base and 3 on rostrum in juv., tending in adult to become
open grooves. Parietal callus in adult nodular, columella concave, anteriorly
crenulate and subcarinate, glaze in adult extending over nearly half the last
whorl, thickened and forming a flat polished ‘sole’, edge free and varicoid.
Outer lip thickened in adult, internally plicate. 25 x 19 mm., and 27 x
19 mm.
Operculum broadly subtriangular, broader than long, serrate on both
margins, 5 X 6 mm. in 23 mm. shell.
Yellowish-grey, outer lip and columella glaze white, aperture within
brownish or violaceous with a pale spiral band which is not visible externally
except faintly in juveniles.
Il4 ANNALS OF THE SOUTH AFRICAN MUSEUM
Radula with c. 65 rows, central plate with 16 cusps, outermost one on
either side minute, intermediate plate oval, lateral plate without denticles
between the 2 cusps.
Fossil, Pleistocene: Durban (Krige, Tomlin).
Dead: Port Elizabeth (Sowerby).
Living: Natal (Krauss); Durban, Delagoa Bay, Mozambique Island
(S. Afr. Mus.). Inhambane, Portuguese East Africa (U.C.T.).
Distribution. Mauritius, Réunion, Seychelles, Madagascar, Red Sea, Aden,
Indo-Pacific.
Nassa gemmulata (Lam.) -
Fig. 22(g)
1816. Lamarck. Tabl. Encyel., Livr. iv, pl. 394, figs. 5 a, 6, and Liste, p. 1 (Nassa clathrata,
n. et f. only). (Not Livr. i, 1827, as given in Tomlin.)
1822. id. Anim. sans. Vert., vil, p. 271 (Buccinum g.).
1859. Chenu. Man. Conchyl., i, fig. 765.
1886. Watson. Challenger Rep., xv, p. 176.
1go1. Melvill & Standen. Proc. Zool. Soc. Lond., 1, p. 412.
1928. Tomlin. loc. cit., p. 318.
Protoconch ? Postnatal whorls 7; axial ribs 14 on 1st whorl, 15 on 2nd,
17 on 3rd, 22 on 4th, 28 on 5th, 24 on 6th, and 20 on last whorl, suture to
suture, straight, oblique (retractive), continued across base; crossed by 3
spiral sulci on 1st—3rd whorls, 4 on 4th—6th, and 5 on later part of 6th and on
7th whorl, sulci narrow and deep especially on later whorls, dividing the ribs
into rounded nodules, the uppermost forming a coronet over the sunken suture;
3 additional sulci on base, and a deep groove separating lowermost nodules
from the reflexed rostrum, which has 4-5 striae. Internal parietal callus
nodiform, columella more or less granulose, glaze extending over half base,
edge free on rostrum. Outer lip internally plicate. 28 x 13:5 mm.
Operculum broader than long, serrate on both margins, 4 x 6 mm. in
28 mm. shell.
Buff with orange or brown irregular suffusions, aperture and glaze white.
Radula with 70 rows, central plate with 9 cusps, intermediate plate oval,
lateral plate with rather stout inner cusp, no denticles between the 2 cusps.
Durban (Smith, ? dead).
Living: Inhambane, Portuguese East Africa (U.C.T.).
Distribution. Red Sea, Persian Gulf, Karachi, Indo-Pacific to Japan.
Nassa plicatella A. Ad.
1852. A. Adams. Proc. Zool. Soc. Lond. (for 1851), p. 111.
1903. Von Martens. D. Tiefsee Exp., vii, p. 9.
1912. Dautzenberg. Ann. Inst. ocean., vol. 5, fasc. 3, p. 32.
1923. Odhner. Géteb. K. Vet. Handl., xxvi, 7, p. 14. .
1928. ‘Tomlin. loc. cit., pp. 323 and 327.
1931. Lamy. Bull. Mus. Paris, (2) III, p. 304.
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), pp. 34, 46.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA P15
Protoconch 14 (2) whorls, diam. 1, alt. 0-6 mm., smooth. Postnatal
whorls 5, profile evenly convex; axial ribs on 1st whorl 12 or 13 (junction with
protoconch indistinct), 13 on 2nd whorl, increasing to 17-18 on 4th and to
20-24 on 5th whorl, suture to suture, slightly oblique (retractive), extending
to base, sometimes duplicated and irregular or obscure on last half whorl;
crossed by spiral lirae 5-6 on 2nd, increasing to 8—g on 5th whorl, intersections
slightly nodulose, 7-8 additional lirae on base, one (? more) on rostrum. No
parietal callus, columella smooth, rather conspicuously carinate anteriorly,
glaze narrow. Outer lip not toothed at either end, internally feebly plicate if
at all. 27 (protoconch missing) X 15 mm.
Operculum triangularly oval, serrate on both margins.
Cream or greyish, darker in the intervals between the ribs when wet,
uniform when dry.
Radula with 70-75 rows, central plate with 10 cusps, the outermost on
either side minute, no intermediate plate, lateral plate without denticles
between the 2 cusps.
Fossil (late Tertiary): Saldanha Bay (Tomlin, Haughton).
Dead: Walfish (“Wallwich’) Bay (Adams, Lamy); Great Fish Bay, Angola
(von Martens); Angra Pequena (Liideritzbucht) (von Martens); Table Bay
(Tomlin, S. Afr. Mus.).
Living: Mossamedes and Praya Amelia, littoral and 15-35 metres
(Dautzenberg); Port Alexander (Odhner); Langebaan, Saldanha Bay, low
tide (U:C.T.).
Remarks. Sowerby’s record from Natal is not acceptable.
The number of ribs varies somewhat: in one specimen there are ?12 on
1st whorl (worn), 13 on 2nd, 12 on grd, 12 on 4th and 13 on 5th whorl, the
intervals consequently being noticeably wider than normal. The Saldanha
Bay examples are transitional.
Walhmerm( 92a) Goieb. Wh. Very Handi, xxv, 7, p. 14, pl.\1, figs. 6, 7)
described angolensis similar to plicatella but smaller, 14:2 x 7-6 mm., also from
Port Alexander, 16 fathoms.
Nassa desmoulioides Sow.
Figs. 22(d), 23(c)
1903. Sowerby. Mar. Invest. S. Afr., 11, p. 219, pl. 4, fig. 1.
1928. Tomlin. loc. cit., p. 317.
1956. Knudsen. Aélantide Rep., 4, p. 49, pl. 2, fig. 3 (laps. cal. desmouleoides).
1957. Franca. Anais 7. Invest. Ultramar., x, 2, p. 29, pl. 1, figs. 1, 2, pl. 2 (laps. cal. desmou-
leoides).
Protoconch 2 (24) whorls, diam. 1-1-25, alt. 0-75-0-8 mm., smooth.
Postnatal whorls 6, profile rounded but with squarish shoulder at the level of
the sunken suture; axial] ribs 16-17 on Ist whorl, increasing to 23-25 on 5th
and 30-32 on 6th whorl, suture to suture, straight, narrower than intervals,
sometimes on later part of 6th whorl becoming irregular or even obsolete;
116 ANNALS OF THE SOUTH AFRICAN MUSEUM
crossed by spiral lirae 7 on 1st and 2nd whorls, 8 on 3rd, increasing to 10-11
on 6th, intersections more or less nodular, especially on upper part of whorls;
7-8 additional lirae on base and 3—4 on rostrum. Parietal callus subcariniform,
columella concave, feebly crenulate and carinate anteriorly, glaze thin,
extending scarcely halfway across base, edge free. Outer lip internally plicate.
21 X 13 mm.
Operculum thin, triangularly ovate, serrate on both margins, 5 X 3 mm.
in 18 mm. shell.
White with irregular blotches and axial streaks of orange-brown, oper-
culum pale corneous.
Radula with 70-80 rows, central plate with 10-12 cusps, the outermost
one on either side minute, no intermediate plate, lateral plate without denticles
between the 2 cusps.
Dead: Zululand to Algoa Bay and Agulhas Bank, 40-100 fathoms: most
northerly locality off Cape Vidal (Zululand), most westerly and southerly
Brown’s Bank, approx. 364° S. 21° E. (S. Afr. Mus. P.F. coll.).
Living: off Cape Natal (Durban), 54 fathoms; off Great Fish Point, 53
fathoms (S. Afr. Mus. P.F. coll.) ; 29° 46’ S. 31° 17’ E. 60-70 fathoms (U.C.T.).
Distribution. Off Sierra Leone (4° 24’-13° 43’ N. 7°-17° 23’ W.), 65-90
metres; off Portuguese Congo (5°S. 11° 14’ E., 55 metres) (Knudsen); off
Cabinda (Franca). :
Remarks. The majority of examples do not quite conform with Sowerby’s
description. He said the ribs number ‘10 on the penultimate [i.e. 5th] whorl’;
this might be correct if he counted only those visible on one half of the whorl,
but his figure showed 6 (or 7). I have seen one 5-whorled specimen and one
juvenile taken together with a normal (as here described) specimen (off Cape
Natal, 54 fathoms) with 12 axial ribs on 1st whorl, 13 on and, 14 on grd, 16 on
4th and 18 on 5th whorl; the intervals between the ribs are consequently very
obviously wider than the ribs. There is no doubt that they are conspecific,
apart from there being an occasional intergrading example.
The occurrence of this species in a locality in the western Indian Ocean
would not have been unexpected, but its discovery by the Atlantzde in tropical
West African waters is very surprising. ‘The West African specimens have been
compared by Knudsen with Sowerby’s type material in the British Museum,
and the identity may thus be accepted.
Nassa speciosa A. Ad.
Fig. 22(c)
1852. A. Adams. Proc. Zool. Soc. Lond. (for 1851), p. 100.
1903. Von Martens. D. Tiefsee Exp., vii, p. 26 (plicosa).
1907-8.* Melvill & Standen. Tr. Roy. Soc. Edinb., xlvi, p. 153 (Phos plicosa).
* Issued separately Sept. 1907. The 1909 Report is the same as that in the Transactions,
with different pagination; the original pagination is given at the foot of each page.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA il7
1909. id. Sci. Res. Scotia, v. p. 123 (Phos plicosa).
1928. Tomlin. loc. cit., p. 324 and p. 327 (plicosus Dnkr., non Menke).
1932. Turton. Mar. Sh. Pt. Alfred, p. 59 (plicosa).
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), pp. 33, 46 (plicosa).
1936. Peile. Proc. Mal. Soc., xxii, p. 140 (plicosa, radula mentioned).
Protoconch 2 (24) whorls, diam. 0:75, alt. 0-5 mm., smooth. Postnatal
whorls 8, profile nearly straight but with a sloping shoulder near top of whorl;
axial ribs 12 (13) on 1st whorl, 12-13 on 2nd, 14-15 on 3rd, 4th, and 5th,
13-14 on 6th, 12-13 on 7th, and 10-12 on last whorl, suture to suture, extending
across base, straight or slightly sigmoid and protractive on base of last whorl,
often (in adult) additional plicae on back of outer lip but not reaching to the
shoulder; crossed by spiral lirae 4-5 on 1st and 2nd whorls, 6 on 3rd, 7 on 4th,
increasing to 12-15 on last whorl, the 3-4 (5) next the suture finer than the
others which are variable, often wider and narrower lirae alternating, or a
wide lira is divided by a fine stria; 8-9 additional lirae on base with finer
intermediaries, 7-8 on rostrum, intersections with ribs on lower half of base
and back of outer lip more or less nodulose. Internal parietal callus blunt,
columella smooth, bluntly carinate anteriorly, glaze extending more than half-
way across base, not very thick, edge adnate except on rostrum. Outer lip
internally more or less plicate. 31 X 16 mm. Two examples from the same
hanlige < 16 mm. and 29°5 xX 14:5 mm.
Operculum oval, margins entire, 8 X 4:5 mm. in 29 mm. shell.
Buff or cream, unicolorous, periostracum brown, columella glaze and
aperture white, anterior canal purplish-brown, operculum dark brown.
Animal pale, speckled with grey.
Radula with c. 80 rows, central plate with antero-lateral angles somewhat
acutely produced, with 9 cusps, no intermediate plate, inner cusp of lateral
plate with 1-3 minute denticles on its outer margin (i.e. facing the outer
cusp).
Fossil, late Tertiary: Saldanha Bay. (Tomlin, Haughton).
Living (and dead): from off Umhloti River (Natal), East London, Port
Alfred, Algoa Bay, Agulhas Bank, False Bay, Table Bay, to Saldanha Bay,
low tide to 50 fathoms (S. Afr. Mus. P.F. coll.); 34° S. 25° 46’ E., 41 fathoms,
Bees. 20. 4 E., 26 fathoms, 33° 3’ S. 28° 11’ E.., 91 fathoms, and 31° 38’ S.
29° 34. E., 26 fathoms (U.C.T.).
Remarks. Plump and slender forms occur. The number of axial ribs
varies slightly. In beach-worn examples the shoulders of the ribs often appear
“as prominent white tubercles.
Von Martens stated that the margins of the operculum were serrate, but
this conflicts with all the specimens in 8S. Afr. Mus., except in one specimen
where the outer edge is crenulate, probably due to wear or corrosion.
Bisacchi (1930. Ann. Mus. Civ. Genoa, lv, p. 59) refers Red Sea examples
to a South African variety figured, but not described, by Marrat (1877. New
Forms .. . Nassa, pl. 1, fig. 11). This is probably a misidentification.
118 ANNALS OF THE SOUTH AFRICAN MUSEUM —
Nassa plebecula Gould
1860. Gould. Proc. Boston Soc. Nat. Hist., vii, p. 332.
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 6, pl. 8, figs. 4, 5 (producta).
1928. Tomlin. loc. cit., p. 323.
1932. Turton. Mar. Sh. Pt. Alfred, p. 57, and p. 57, pl. 13, no. 424 (subcancellata).
? 1932. id. ibid., p. 57, pl. 13, no. 423 (erecta).
Four worn specimens in 8. Afr. Mus., with apices worn and only 5 or 6
whorls remaining. Sowerby gave the number (total) of whorls as 9, which
seems a little excessive even if the protoconch were included.
On the last 4 remaining whorls the axial ribs increase regularly by one rib
on each successive whorl, the numbers being resp. 11--13, 12-14, 13-15, and
14-16. The spiral lirae number resp, 4, 5, 6, and 6; the uppermost lira imme-
diately below the suture of preceding whorl is slender, and on last whorl or
2 whorls a 7th lira may be partly visible in the suture with the following whorl.
5-6 additional lirae on base.
Sowerby said the outer lip was plicate within, and the columella rugose;
his figure shows the plicae and a small parietal nodule. ‘The present specimens
show no parietal nodule, or labral tooth, or plications, the columella is smooth,
and the glaze narrow.
13-5 X 5 mm. (Sowerby); 12 X 4:5 mm. (S. Afr. Mus) iunbam
(Sowerby).
Distribution. Mauritius (Tomlin), Japan (Gould), Polynesia (Tomlin).
Remarks. Tomlin said the type of producta (in British Museum) was a poor
specimen, but Sowerby’s figure gives the impression of a rather well-preserved
specimen.
Nassa papillosa (Linn.)
1758. Linne. Syst. Nat., 1oth ed., p. 737, sp. 393 (Buccinum p.).
1816. Lamarck. Tabl. Encycl. Meth., pl. 400, figs. 2 a, b, and Liste, p. 2.
1880. Von Martens. Mauritius G Seychellen, p. 241.
1952. Braga. Anais Est. Zool. Ultramar., vii, 3, p. 75, pl. 2, fig. 5.
Braga records this species (dead) from Mozambique. It occurs in
Mauritius, Réunion, Madagascar, Aden, and Indo-Pacific.
Nassa vidalensis n. sp.
Fig. 24(d)
Protoconch 3% whorls, diam. 1-3, alt. 1:2 mm., smooth, and and 3rd
whorls with peripheral keel, which sinks down on last half whorl into the
suture with 1st postnatal whorl, junction with ist postnatal whorl deeply
concave. Postnatal whorls 3, slightly turreted, sutures deep, profile gently
convex; all whorls smooth, without any axial or spiral sculpture, except faint
indications of 2—3 fine spiral striae at top of 2nd and 3rd whorls, best seen at
back of outer lip, indications of 4—5 striae at bottom of base. Outer lip thickened,
varicoid, feebly plicate within. 6 X 4 mm.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 119
Off Cape Vidal (Zululand), 80-100 fathoms (S. Afr. Mus. no. A88309,
EP coll.).
Remarks. A single unworn specimen, similar in shape to dissimilis Watson
1886, and aracanensis Smith 1899, 1901; and with carinate protoconch similar
to that of babylonica Watson 1882, 1886, agapeta Watson 1882, 1886, and psila
Watson 1882, 1886.
1848.
1906.
1920.
1928.
1936.
1952.
Fic. 24.
(a) tNassa scopularcus n. sp. (b) N. vidalensisn. sp. (c) Nassa sp. juv.
Nassa kraussiana (Dnukr.)
Figs. 22(k), 23( f )
Krauss. Siidafr. Moll., p. 123, pl. 6, fig. 18.
Smith in Rogers. t1oth Rep. Geol. Comm. (Cape) for 1905, p. 291.
Wybergh. Tr. Geol. Soc. S. Afr., xxii (1919), p. 66.
Tomlin. loc. cit., p. 320 (references), and p. 327.
Peile. Proc. Mal. Soc., xxii, p. 140, fig. 4 (radula).
Braga. Anais Est. Zool. Ultramar., vii, 3, p. 75, pl. 3, fig. 3.
Broadly oval when young, obliquely ovoid and plano-convex when adult.
Protoconch 14 whorls, diam. 0-5, alt. 0-3 mm., smooth. Postnatal whorls 4;
axial ribs 14-15 on Ist whorl, 15-16 on 2nd, c. 16 on 3rd but feeble and evanes-
120 ANNALS OF THE SOUTH AFRICAN MUSEUM
cent on later part of whorl, none on 4th whorl, but usually a few irregular
plicae on back of outer lip, ribs extending from suture to suture, slightly
oblique (retractive) ; 4—5 striae on back of outer lip anteriorly (not visible after
callus is fully formed); sometimes a well-incised spiral stria in upper part of
Ath whorl (distant about 7 of height of whorl from suture) traceable back on
to the 3rd and part of 2nd whorl where it is lost among the axial ribs; sometimes
also 3—5 similar striae below the periphery but less distinct than those on base.
Columella concave, smooth; glaze thick and extensive, when fully developed
spreading over whole of base and enveloping half the shell up to apex, and
forming with the callously thickened outer lip a smooth polished ‘sole’, the
actual aperture occupying only about $ or @ part of the sole. 11 X 8 mm.
13 mm. long (Braga). Smallest example with fully developed callus
6 X 4°75 mm.
Operculum broader than long, almost semicircular, apex worn, one or
two strong teeth on both margins.
Ochraceous with 3 purplish-brown spiral bands (sutural, peripheral, and
basal), ‘sole’ white, yellowish, or (especially outer lip) purplish; back often
stained green with algae.
Radula with 55-65 rows, central plate with 12 cusps and a minute one
on either side externally, no intermediate plate, lateral plate with inner cusp
broad, curving inwards (medianly), its inner (median) margin with 3-5
denticles.
Fossil (late ‘Tertiary, Pleistocene, and Recent): Bredasdorp (Wybergh) ;
Durban, Port Elizabeth, Plettenberg Bay, Knysna, Little Brak River (see
Tomlin, p. 327). Also Sedgefield, near Knysna (A. R. H. Martin. S. Afr. 7.
Sci., 52, p. 187, 1956).
Dead: Durban, Port Alfred, Port Elizabeth, Jeffreys Bay, Still Bay (auct.
and 8S. Afr. Mus.).
Living: Keurbooms River estuary (Plettenberg Bay) ; Delagoa Bay (S. Afr.
Mus. coll. K.H.B.). Delagoa Bay (U.W.). Inhambane, Portuguese East
Africa (U.C.T.).
Remarks. ‘This characteristic species is an estuarine and littoral species;
the specimens taken in 43 fathoms off East London (recorded by Tomlin) were
dead.
Peile (1936, loc. cit., and 1939, Proc. Mal. Soc., xxiii, p. 276) says sordidus,
echinatus, creniliratus and thersites have similar lateral plates in the radula, the
last two with intermediate plates in addition.
tNassa scopularcus n. sp.
Fig. 24(a)
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 20 (cf. babylonicus).
Turreted, resembling in general babylonica Watson, with strong axial ribs
and, at first sight, no spiral sculpture.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA E20
Whorls 54 (6) including protoconch (tip of which is missing); axial ribs
on 2nd whorl worn, not countable with certainty, 11 on 3rd and 4th, 12 on
last whorl, from suture to suture, extending over base to columella glaze,
straight, their tops forming a squarish subsutural shoulder. Faint traces of
spiral lirae (? 6) between the ribs on part of 4th whorl, and 13 more distinct
on last portion of last whorl. Outer lip not plicate within. 14 x 7°3 mm.
White.
Late Tertiary marine beds, shore of lagoon, south of Bogenfels, South
West Africa (S. Afr. Mus. Ag230 coll. S. H. Haughton).
This shell was seen by Tomlin, who said ‘nothing Recent like it except
babylonica, and it is quite distinct from that’. It has fewer axial ribs; and when
fresh the spiral lirae would probably have been much more prominent than in
babylonica.
Bogenfels = Arch-rock = scopularcus.
Nassa sp. juv.
Fig. 24(c)
Juv.—Protoconch 5 whorls, alt. and diam. 1-3 mm., smooth, profile of
the last whorl strongly convex, not carinate, a few pliculae before the junction
with Ist postnatal whorl, which is abrupt. Postnatal whorls 2, shouldered;
axial ribs 13 on each whorl, forming small points at the shoulder, slightly
retractive; crossed by spiral lirae on 1st whorl 1 between suture and shoulder,
later 2, on 2nd whorl 2, below shoulder 3 on each whorl, on 2nd whorl one
intermediary between each pair, on base 3 additional and 2 on rostrum, with
intermediaries. 5°5 X 3 mm.
Protoconch and rst whorl white, 2nd whorl! pale buff, faintly brown above
shoulder, and with 2 faint brown bands below, one from upper margin of
aperture, the other from about middle of columella.
Off Umkomaas (Natal), 40 fathoms, one (S. Afr. Mus., Reg. No. A8888,
P.F. coll.).
Remarks. Larger, 5-5 mm. with 2 (postnatal) whorls, than the East Indies
patricia Thiele 1925, 3°5 with 3 whorls; but otherwise similar, and likewise with
two brown bands. Thiele said the protoconch of patricia was feebly keeled on
last whorl (as in babylonica), but in the present specimen it is not keeled and
moreover is very much more prominent; in fact much like that of some
Cymatiids or Bursids.
The resemblance to arakanensis Smith 1899 and 1901 is not so close, and
here also the protoconch (especially the 5th whorl) is not nearly so prominent,
and the spiral lirae are more numerous.
Although quite distinct among the South African species, this juvenile
may belong to one of the numerous Indo-Pacific species, and is better left pro
tem. without a name.
122 ANNALS OF THE SOUTH AFRICAN MUSEUM
Gen. DEmouLIA Gray
1838. Gray. Ann. Mag. Nat. Hist., i, p. 29.
1847. id., Proc. Zool. Soc. Lond., p. 140 (Desmoulea).
1926. Tombln.” Am S.A, as, soy, p- 326.
1929. Thiele. Handbuch, i, p. 323 (Desmoulea, subgen. of Nassa).
Demoulia abbreviata (Gmelin)
Fig. 22(/)
1928. Tomlin. loc. cit., p. 326.
1932. Turton. Mar. Sh. Pt. Alfred, p. 60, pl. 14, no. 441 (juv.).
Subglobular, with numerous conspicuous spiral costae. Protoconch 2
whorls (tip missing), diam. 1:25-1°5 mm., smooth. Postnatal whorls 6 (7).
Periostracum thin, fibrous-fimbriate. 38 (protoconch missing) x 28 mm.;
plump and slender forms 33 x 26 mm. and 35 xX 24 mm.
Operculum thin, triangular, edges scarious, 9 X 9 mm. in 30 mm. shell.
White, periostracum brown, operculum amber.
Radula with c. 95 rows, central plate with 9 cusps, lateral plate with inner
cusp slender.
An egg-capsule removed from oviduct of animal in 30 mm. shell was
13 mm. in length, and 3-3-5 mm. in cross-section, cylindrical, somewhat
triquetral in section (but this may not be the true shape when the capsule
has been laid and has hardened).
False Bay to Algoa Bay and Natal, 18-52 fathoms (Tomlin, S. Afr. Mus.
P.F. coll.). St. Francis Bay, 80-100 metres (von Martens).
Living: False Bay, 27 fathoms; off Knysna, 52 fathoms; Algoa Bay,
31 fathoms (S. Afr. Mus. P.F. coll.).
Demoulia retusa (Lam.)
1816. Lamarck. Tabl. Encycl. Meth. Livr., iv, pl. 394, figs. 3 a, b, and Liste, p. 1. (Nassa
ventricosa n. et f. only) (not Livr. iii, 1827, as given by Tomlin).)
1822. id. Anim. sans Vert., vii, p. 270 (Buccinum r.).
1926; Vonlin. \loc: cit., "9.926.
Ovoid, with numerous fine spiral lirae. Protoconch 2? whorls (tip missing),
diam. 1:3 mm., smooth. Postnaial whorls 5 (6). Periostracum thin, smooth.
27 (protoconch missing) x 19 mm.
Variously blotched and marked with brown, usually two speckled spiral
bands and ‘necklace’ of dark and pale spots at top of whorls, periostracum pale
brown.
False Bay, Agulhas Bank, and Zululand, 5-55 fathoms (Tomlin, S. Afr.
Mus. P.F. coll.). 7
Three living specimens were obtained by the Pieter Faure in False Bay,
5 fathoms, but the animals were not preserved.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 123
Gen. Buus Griffith
1834. Griffith in Cuv. Anim. Kingd., xii, pl. 37.
1840. Swainson. Treat Malac., p. 302 (Leiodomus).
1841. Gray. Proc. Zool. Soc. Lond., xv, p. 139 (Dorsanum).
1929. Thiele. Handbuch, i, p. 322.
1937. Peile. Proc. Mal. Soc., xxii, p. 183 (radulae) (also comments on Thiele’s arrangement).
1938. id. ibid., xxiii, p. 6 (radula).
Protoconch, junction with Ist postnatal whorl not sharply demarcated.
Operculum with apical nucleus, but shape variable: of more or less normal
size, subtriangular, margins entire or serrate; or ovate with incurved apex;
or much reduced in size.
Animal with large foot capable of considerable expansion, its antero-
lateral corner more or less pointed (tentaculate) (Bullia) or rounded (Dorsanum),
posteriorly with two ‘tails’ (Bullia) or without tails (Dorsanum); eyes present
(Dorsanum) or absent (Bullia).
Radula formula 1.1.1, central plate with several cusps, lateral plate with
2 large cusps, the inner one bifurcate (Bullia) or trifurcate (Lezodomus) or with
additional denticles between it and the outer cusp. Considerable variation
occurs in the cusps of the lateral plates, even in the same radula (cf. Peile,
1937), and this fact is somewhat against Peile’s argument for separating
Leiodomus from Bullia (loc. cit., p. 184).
The radulae of eleven species, including those already described by
Peile, are here described and nine of them are figured.
Remarks. Many parts of the South African coast, where sandy beaches
occur, await exploration, and doubtless wil) yield interesting results. Needless
to say: living animals are required for study.
With the exception of the egg-capsule of tenuis described below, the
reproduction and life-history of these arenicolous molluscs seem to be unknown.
I have found juveniles of rhodostoma as small as 4 mm. long in the shifting sand
between tide marks. Do the adults (of the littoral species) retreat off-shore to less
turbulent water for spawning, or do they burrow deeply between tide marks?
Ancilla osculata Sow. is transferred to the present genus, and the suggestion
is made (see also p. 62) that Bullza ancillaeformis Smith is really an Ancilla.
Key based on the operculum
» 1. Operculum of more or less normal size.
A. Margins serrate.
Hes We WCRI CCeAL SUILULES, 19 (oom yet) Got hess) ik vay aye SUMIELES
Pea Slie llamo ChNIPECs "ss ea ce) hea wo ec aoe ke. wo eettalas
B. Margins smooth.
i oubiriane ilar, apex truncate or excavateys i) 2 1s. ).2 acs
annulata
2. Ovate, apex subacute, incurved.
ds Almesticompletely filling aperture yey 8 natalensis
b. Not completely filling aperture.
Pe OMeMEsIMOOE "rue eae ees fg 5.) ) FROdOStOMA
Tem OMC UMA tris ls ecg omuinviem te A es oe ts. ws Ue
Mey Shell pUStULOSE iy. 2) sls tegeMinenes bas ue. “eo iba a wmmozambicensis
124. ANNALS OF THE SOUTH AFRICAN MUSEUM
II. Operculum much reduced in size.
A. Aperture of shell longer than spire. .. .....450 =... = = eee
B. Aperture subequal to spire. Parietal callus thick . . . . . . callosa
C. Aperture less than spire (adult). Parietal callus thin . . . . . tenuis
Operculum unknown: diluta, trifasciata, tenuistriata, ancillaeformis,* dulcis Sow.,
(= digitalis), almo Bartsch (= digitalis), lara Bartsch (= ? tenuis), aepynota
Bartsch (= ? diluta), alfredensis Bartsch (= diluta); and all Turton’s ‘species’.
Bullia similis Sow.
Figs. 25(2), (J), 27(@)
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 5, pl. 7, fig. 1.
Aperture 1-14 times in spire. Protoconch diam. 0-3-0-5 mm. Whorls
(total) 8 (84). Parietal callus thin and narrow, not extending on to preceding
whorl or visible at suture. Whorls smooth with a spiral sulcus near suture, the
lira thus demarcated producing with the growth-lines a crimped or beaded
sculpture beginning (clearly visible) from about 4th whorl; 6th whorl with
1-3 (4) very fine spiral striae, increasing to 6-7 on 7th whorl and 8-9 (some-
times 10-11) on last whorl, becoming stronger and easily visible on the latter,
especially on upper and lower parts of whorl; on last whorl the crimping
extends on to the lira below the sutural lira; growth-lines fine and close,
making the striae punctate, especially on base; 7-8 additional striae on base,
becoming stronger (sulci) near rostrum. Columella glaze thin. 26 x 11 mm.
31 X 15 mm. (Sowerby).
Operculum thin, triangularly ovoid, both margins serrate, 6 X 3°5 mm.
in 21 mm. shell with 9 mm. aperture.
Cream or buff, with very faint orange or fawn undulate, arcuate, or
zigzag axial flames.
Radula with about 60 rows, central plate subequal in width to lateral
plate, base (excl. cusps) about 24 times as wide as long, with 11 cusps, some-
times a minute denticle externally on one side or the other, lateral plate with
inner cusp bifurcate, outer prong a little smaller and more slender than the
inner (median) prong, sometimes a minute denticle between the prongs, outer
prong itself sometimes bifid or bifurcate.
Fossil: raised beach, alt. 375 ft. at Durban-Umgeni (Geol. Surv.).
Living: off Cone Point (between St. Lucia and Durnford Bays, Zululand)
to off East London, 12-27 fathoms (S. Afr. Mus. P.F. coll.).
Remarks. Close to belangeri Kien. in coloration, but the latter, as already
remarked by Sowerby, has no crimping near the suture; also (Ceylanese
specimens in S. Afr. Mus.) the middle portion of the whorls is without spiral
striae.
* See note under Ancilla bullioides, p. 62.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA P25
Fic. 25.
Central and lateral radula plates of Bullia (a) pura Melv.; (6) tenuis Gray; (c) natalensis (Krss.) ;
(d) mozambicensis Smith; (e) digitalis Meuschen; (jf) annulata (Lam.); (g) rhodostoma Rve.;
(hk) laevissima (Gmelin), with variants of lateral plate; (2), (j) similis Sow., with variants of
inner cusp of lateral plate. In (j) the right-hand one is from the same radula as figured above;
the three on the left are all from another radula.
126 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bullia osculata (Sow.)
Fig. 27(b)
1900. Sowerby. Proc. Mal. Soc., iv, p. 3, pl. 1, fig. 6 (Ancilla o.).
1903. Smith. ibid., v, p. 364 (Ancilla o.).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 35 (Ancilla o.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 32, pl. 7, no. 243 (Ancilla o.).
Aperture 1-1} in spire. Whorls 6-7. Parietal callus thin and narrow, not
extending on to preceding whorl and not visible at sutures. A single shallow
and inconspicuous spiral stria immediately below the suture, and 2-3 on
lower part of base (excluding those on rostrum). Columella glaze thin.
19 x 6 mm, ~Wurton: 21) mm, long:
Operculum triangularly ovoid, margins smooth, apex truncate, 3°5 xX
2 mm. in shell 14 mm. with aperture 5:5 mm.
Cream, with flame-like markings below the suture and around base, in
addition with axial undulate lines (Turton, and S. Afr. Mus.); in the Cape
Vidal specimens these lines are strongly zigzag.
Foot with antero-lateral corners rounded (but ?, only one example
available), posteriorly with 2 well-developed ‘tails’, no eyes. Radula very like
that of semilis, with 55 rows, central plate about twice as wide as long (excl.
cusps), with 11 cusps, the outermost one on one side minute, lateral plate sub-
equal in width to central, inner cusp with both prongs stout, the inner larger
than the outer.
Living: off Cove Rock (East London area), 22 fathoms (S. Afr. Mus.
PF. coll.)
Dead: Pondoland, the Kowie, Port Alfred (Sowerby, Bartsch, Turton).
Cape Vidal, Zululand, 50 fathoms (S. Afr. Mus. P.F. coll.).
Remarks. ‘Vopotypes in 8. Afr. Museum. The Cape Vidal specimens were
identified by Sowerby, but not recorded.
Sowerby commented on the likeness of his species to a Bullia; indeed it
seems surprising that he referred it to Ancilla because it lacks the basal groove
characteristic of the Olividae. Neither Smith nor Bartsch queried its generic
position. I had previously transferred it to Bullia on conchological grounds,
but now a specimen containing the animal, found in one of the P.F. bottom-
samples, settles the question.
Sowerby distinguished similis from belangert Kien., but the present species
is much nearer to the latter. Two Ceylanese specimens in S. Afr. Mus. (ex
coll. Ross-Frames), 35 mm. long, have axial lines and flames, but they have
two spiral striae at the top of each whorl, distinct from the early whorls onwards,
the upper one of which becomes a sulcus on the later whorls; there are also 2
striae on lower part of the whorl.
The species is very like a slender form of annulata (of equal size and before
the typical shoulder of the later whorls has developed), especially smooth
examples in which the spiral striae are inconspicuous and obsolescent.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 297)
Bullia tenuistriata Tomlin
1920. Tomlin. 7. Conch., xvi, p. 87, text-fig. 4.
1923. id. ibid., xvii, p. 46.
1932. Turton. Mar. Sh. Pt. Alfred, p. 64.
Aperture about 14-14 times in spire (as figured). Sutures deeply impressed ;
presumably therefore the parietal callus is thin and not visible on the preceding
whorl. 17 spiral striae on penultimate whorl, and 15 on each of the preceding
whorls, interstices smooth. Otherwise similar to pura. 16°5 X 65 mm.
Turton: 25 X 10 mm.
Port Alfred.
Described from two dead specimens. Turton later found a larger one.
Remarks. ‘Tomlin said it had ‘some slight resemblance’ to pura. In my
opinion it has a very strong resemblance, amounting almost to identity. Com-
pared with some specimens of pura in which the intermediary lirae are well
developed (total number of lirae 15-17!) the only distinguishing feature is the
sunken suture of tenuistriata, and that may well be only an individual aberration
(cf. alfredensis Bartsch and scalaris Turton, both = diluta).
Bulla trifasciata Smith
Fig. 27(f)
1go4. smith. 7. Malac., xi, p. 34, pl. 2, fig. 17.
Distinguished from annulata, apart from the coloration, by being spirally
striate from the 2nd whorl onwards, and by the uppermost stria (juv.) or
(older specimens) the 2 uppermost striae being on the actual turgid shoulder.
Apex blunter, 1st whorl diam. at least 1-5 mm. (annulata: 1 mm.). Whorls
(total) 7, turreted or pagoda-like, profile below shoulder almost straight (at
least in later whorls). In perfectly fresh specimens the spiral striae could
probably be traced on the 1st whorl (I have seen only worn specimens); on
2nd whorl 7 striae, on later whorls 8, 7-8 additional ones on base. 32 X 15 mm.
Smith: 39 x 18 mm.
Operculum and animal unknown.
Brown with 3 darker spiral bands, that on the shoulder being the FEV eee:
bleached specimens white with the bands more or less visible.
Dead: Port Alfred (Smith, Bartsch, Turton); Still Bay (S. Afr. Mus. coll.
Muir).
_ Remarks. Smith did not mention the one essential difference in the spiral
sculpture between this species and annulata.
Bullia annulata (Lam.) Rve.
Figs. 25(f), 27(¢)
1816. Lamarck. Tabl. Encycl. Meth., p. 399, figs. 4.a, 6, Liste, p. 2 (Buccinum a., nom. et fig.).
1846. Reeve. Conch. Icon., iii, Bullia no. 13.
1902. Sowerby. Mar. Invest. S. Afr., ii, p. 95, pl. 2, fig. 4 (with animal).
1903. Von Martens. D. Tiefsee Exp., vii, pp. 28, 53.
128 ANNALS OF THE SOUTH AFRICAN MUSEUM
1913. Bullen Newton. Rec. Albany Mus., ii, p. 344, pl. 23, figs. 3, 4.
1937. Peile. loc. cit., pp. 183, 184, fig. 17 (radula).
Aperture in juv. subequal to spire, later 1g-14 in spire. Apical whorl
diam. o-75-1 mm. Whorls (total) 9 (94), turreted or pagoda-like, profile
below shoulder slightly convex. Parietal callus thin, not extending on to
preceding whorl nor visible at suture on early whorls, but on later whorls often
visible as a narrow irregularly undulate excrescence between suture and
preceding whorl (sometimes visible and invisible on different parts of the same
shell). No sculpture on first 5 whorls except from 2nd whorl onwards a single
spiral stria on upper part of whorl near suture; on 6th and later whorls the
subsutural stria becomes a sulcus and is followed by 7-8 striae across the whole
whorl, on last 2 or 3 whorls these striae become stronger and can also be called
sulci; the lira adjoining the suture from the 4th or 5th whorl onwards becomes
stronger and forms a tabulate shoulder varying in strength, sometimes sloping,
sometimes projecting almost perpendicular to the preceding whorl; 9-10
additional striae (sulci in large examples) on base. Anterior end of columella
prominent, angularly carinate; glaze thin. 23 x 12 mm. 36 X Ig mm.,
45 X 22 mm., 60 x 28 mm.; smallest specimen examined 4 X 2°75 mm. (but
see infra). |
Operculum in adult broadly triangularly ovoid, margins entire, apex
truncate; in juveniles obliquely oblong, broader than long, apex excavate
between 2 small points; 10 x 8 mm. in 39 mm. shell with 16 mm. aperture;
juv. 2 X 3 mm. in 13 mm. shell with 6 mm. aperture.
Buff or fawn, with violaceous (darker fawn or fulvous in dead shells)
marks on the shoulder, which frequently, especially in juveniles and half-grown
examples, extend across the whorls as axial streaks and flames; operculum
amber-brown. |
Foot with antero-lateral angles pointed (or tentaculate, depending on the
preservation), posteriorly with 2 long ‘tails’, no eyes. Radula with 85-90 rows,
central plate about 14 times as wide as lateral plate, base (excl. cusps) about
34 times as wide as long, with 12 cusps, lateral plate with inner cusp bifurcate,
its outer prong smaller and more slender than inner (median) prong.
Animal pale greyish, livery, or flesh-coloured.
Fossil, Mio-Plhocene: Redhouse near Rort Elizabeth (Newton).
Port Alfred (Bartsch, Turton); St. Francis Bay, 80-100 metres (von
Martens); off Nieca River (East London area), 43 fathoms, 1 dead (S. Afr.
Mis. PF. colly):
Living: False Bay, 18-20 fathoms; off Cape St. Blaize, 19 fathoms; Algoa
Bay, 10-24 fathoms (S. Afr. Mus. P.F. coll.). Knysna and Algoa Bay (U.C.T.);
34° S. 25° 46’ E., 41 fathoms and 33° 47'S. 26° E., 26 fathoms (U.C‘T.):
The typical form has not been recorded farther east than Port Alfred and
East London area, and farther west than False Bay.
Remarks. Sowerby’s 1go02 figure of the animal shows the two long “tails’,
but not the true shape of the operculum.
8
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 129
Examples 10-11 mm. long with 5 whorls and the beginning of the 6th are
the smallest I have seen which can without doubt be referred to this species,
as they have striae on the middle of the last whorl. Below this size owing to
the absence of these striae juveniles are not separable with certainty from
juveniles of laevissima, although in the latter at an early stage the length of the
aperture exceeds that of the spire.
Peile figured the radula of a juvenile ‘just emerging from the earliest
stages of growth’.
The Algoa Bay specimen recorded by von Martens as mauritiana, a species
closely resembling annulata, is probably a misidentification.
There are 3 typical specimens in S. Afr. Mus. from Zanzibar, collected by
E. L. Layard, on board H.M.S. Castor, 1856. Also a specimen from ‘East
Africa’ (ex coll. Ross-Frames) not so strongly shouldered as typical specimens.
One specimen 23 X 12 mm., 7 whorls, from Still Bay (coll. Muir) is very
smooth. The 6th and 7th whorls each have only 2 striae (one below the
shoulder and one other), only the later part of the 7th whorl has faint traces
of the usual 7-8 striae, and those on lower part of base are only moderately
strong (normally they are strong enough to be called sulci). Although this
specimen is a normal annulata in shape (proportions), the suppression of the
spiral striae connects it with the following form or variety from the Natal coast.
Variety. In general similar to typical form but more slender and less
strongly shouldered, especially on the earlier whorls. The 7th and 8th whorls
show traces of the 2 striae at the top of the whorl, but even on the 8th whorl
the upper one below the shoulder is very shallow and can scarcely be called a
sulcus. In the largest specimen traces of 2 more striae on the upper part, and
one or two on the lower part of the 8th whorl can be seen; and 7-8 additional
ones on*base, the lowermost of which are very shallow sulci.
Operculum as in annulata. Coloration as in annulata.
oe one-time. 35 < 16) (mm... 396 <X116°5 mm., 99 X 18 mm., and
43 X 19 mm. (contrast measurements given above for typical form).
Natal: fresh and somewhat worn specimens off Umkomaas, 13 fathoms;
one fresh specimen off Durban, 54 fathoms; one fresh specimen off Tugela
River, 37 fathoms; one with operculum (presumably taken alive, though
animal not preserved) off Umhloti River, 27 fathoms (S. Afr. Mus. P.F. coll.).
Von Martens (1903) recorded annulata from Natal.
Tomlin saw the Tugela specimen and labelled it ‘annulata var.’.
+Bullia magna Haughton
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 46, pl. 5, figs. 1, 4, 5.
Described from Quaternary deposits 20 miles north of the Orange River
mouth, and from two localities near the Olifants River mouth.
Except for the absence of shoulders the shells might be annulata, and the
anterior end of the columella projects prominently inwards as in this species,
130 ANNALS OF THE SOUTH AFRICAN MUSEUM
but the parietal callus forms a ring ‘behind the posterior margin of each
whorl’, which it does not in annulata.
Moreover annulata has not been recorded from the west coast.
Bullia tenuis Gray
Figs. 25(b), 26, 27(g)
1828. Wood. Suppl. Index Test., p. 12 (nom et fig.).
1839. Gray. ool. of the ‘Blossom’, p. 128.
1903. Von Martens. D. Tiefsee Exp., vu, p. 29.
? 1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 53, pl. 38, fig. 3 (lara).
1932. Turton. Mar. Sh. Pt. Alfred, p. 61 and p. 61 (lara).
Aperture a little longer than spire (juv. 4 mm.), subequal to spire
(c. 12 mm. shells), and a little less than spire (1% times in spire) in larger shells.
Apical whorl (protoconch) diam. 0-8-1 mm. Whorls (total) 8. Parietal callus
Fic. 26.
Bullia tenuis Gray. Egg-capsule with embryo.
Central plate of the 1st, 5th, and 1oth rows of
the radula of this embryo.
on 3rd whorl extending } of whorl
above suture forming a prominent
costa, less prominent on 4th whorl,
and from 5th or 6th onwards forming
only a thin glaze extending +4
length of preceding whorl. On 4th
and following whorls spiral striae
over whole whorl, stronger at top
and bottom, on later whorls (3) 4-5
at top and 4—5 at bottom persist but
striae obsolescent on peripheral area,
the striae at top closer together than
those below, 8 additional striae on
base. Columella glaze thin but
extending over whole base and
rostrum. 60 < 28 mms, ‘smallest
examined (beach-worn) 4 X 2°75
mm.
Operculum very small, oval or
slightly triangularly ovoid, 4:5 x
2°5 mm. in shell 60 mm. with 28 mm.
aperture.
Cream or ochraceous, proto-
conch and apical whorls often with
a livery tinge; operculum amber.
Foot with antero-lateral corners very shortly pointed, posterior tails small,
no eyes. Radula with 75-85 rows, central plate 14 times as wide as lateral
plate, its base 3-34 times as wide as long, with g cusps and a denticle externally
on one or both sides; lateral plate with inner cusp bifurcate, outer prong with
1-3 denticles or serrulations on its outer edge.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA I3I
Animal purplish-grey or flesh-coloured, edge of foot pale.
Four egg-capsules were taken by U.C.T. in False Bay on 29 July 1952.
They are thin and membranous, soft, transparent, with an attachment thread
at each end, one of the threads being less coiled than the other on all four
capsules. One capsule contained numerous eggs, c. 0-2 mm. diam.; the others
only one shell each.
Shell 5:3 X 3°5 mm., aperture 3°3 mm., operculum 0-75 X 0-5 mm.
Radula with 38 rows, central plate of ist row with 3 cusps, of 5th row
with 5 cusps, of roth row with 7 cusps, thereafter from about the 12th row with
g cusps; inner cusp of the lateral plate of the first two or three rows simple,
thereafter bifurcate, sometimes trifurcate.
False Bay to Port Alfred (auct.). Port Natal (Durban) (von Martens,
coll. Heynemann in Berlin Mus.).
Living: Algoa Bay (von Martens); False Bay and Mossel Bay (U.C.T.);
False Bay to off Great Fish Point and Port Alfred, 9-66 fathoms (S. Afr. Mus.
mee ecoll): 39°47 S. 26° E., 26 fathoms, and 33° S. 26° 11’ E., 31 fathoms
(U.G.T.).
Remarks. The ridge-like callus on 3rd and 4th whorls is distinctive.
There are plump and slender forms, e.g. 31 x 16 and 34 X 15 mm.,
47 X 25 and 50 X 23 mm. One specimen is slightly turreted owing to a small
rounded shoulder below the suture on the later whorls.
B. lara Bartsch is probably synonymous, though it is said (by Turton) to
lack the callus ridge, and (as far as can be judged from Bartsch’s figure) the
columellar glaze is not nearly so extensive.
Peansnton (1932. Ir. Geol. Soc..S. Afr., xxxiv (1931), p- 47,pl. 5, fig. 2)
has recorded this species, identified by Tomlin, from Quaternary raised
beaches near the mouths of the Orange River and Olifants River. The figure
shows a shell shaped more like digitalis than tenuis; and on the west coast the
occurrence of tenuis seems rather unlikely.
Bullia pura Melv.
Figs. 25(a), 27(c)
1885. Melvill. 7. Conch., iv, p. 316.
1921. Sowerby. Proc. Mal. Soc., xiv, p. 127 (var. balteata).
1931. Tomlin. Ann. Natal Mus., vi, p. 430.
1932. Turton. Mar. Sh. Pt. Alfred, p. 61, pl. 14, no. 447, and var. balteata, p. 61, pl.14,
no. 448.
1932. id. ibid., p. 61, pl. 14, no. 449 (kraussi).
1937. Peile. loc. cit., p. 184, fig. 18 (radula).
Aperture 14-14 times in spire. Apical whorl (protoconch) diam. 0-5 mm.
Whorls (total) 9 (10); profile of shell from apex to body whorl slightly concave.
Parietal callus thin, not visible above suture except on Jast whorl (or last part
of last whorl), where it extends only a short distance $-+ on to preceding whorl,
but just before and at the aperture it spreads out slightly to } length of preceding
132 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic... 27.
Portions of penultimate and ultimate whorls, and aperture of Bullia to show parietal callus
(stippled) and sculpture. Opercula included when known. (a) similis Sow. (6) osculata (Sow.).
(c) pura Melv. (d) diluta (Krss.). (e) annulata (Lam.) Rve., with operculum of juvenile.
(/ ) trifasciata Smith, posterior part of aperture. (g) tenuis Gray, with 3rd whorl above. (A) natal-
ensis (IKrss.). (2) mozambicensis Smith. (j) digitalis Meuschen. (k) rhodostoma Rve. (1) laevissima
(Gmelin), with variant of operculum. (m) callosa (Gray) Rve.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 1393
whorl above the suture. First 2 (24) whorls smooth; on 3rd and following
whorls 7-9 spiral lirae, on the later whorls often with intermediaries, 5—6 (7)
additional lirae on base, also sometimes with 2-3 intermediaries. Columellar
glaze thin, spreading slightly above posterior canal, covering rostrum but not
more than about 4 of base. 33 X 13 mm., smallest example seen 1°75 XK 1 mm.
(3 whorls). Turton: 35 mm.
Operculum ovate, apex incurved, a slight sigmoid ridge or thickening on
inner surface near inner (left) margin, margins smooth, 4:5 X 2°75 mm. in
shell 22 mm. with 13:5 mm. aperture.
Cream or buff, sometimes with a white band below suture (dalteata), some-
times with irregular faint brown marks around periphery of last whorl or last
two whorls; operculum amber.
Radula with c. 60 (half-grown)—75 rows, central plate about 14 times as
wide as lateral plate, its base 44-5 times as wide as long, with Io rather stout
and bluntish cusps (throughout the radula, not only the anterior ones which
get blunted by wear); lateral plate with inner cusp stout, bifurcate, inner
prong broad with sinuous inner margin, outer prong slender.
Fossil: raised beaches at Keurbooms River estuary (K.H.B. 1931).
Dead: Port Alfred and Port Elizabeth (Sowerby, Bartsch, Turton). Still
Bay (S. Afr. Mus. coll. Muir).
Living: False Bay (S. Afr. Mus.).
Remarks. The tapering spire due to the slenderness of the early whorls
contrasted with the rather disproportionate width of the last whorl, producing
the concave profile of the shell as a whole, is distinctive.
The shape of the inner cusp of the lateral plate of the radula is, so far as
is known, unique among the South African species.
B. tenustriata ‘Tomlin (supra p. 127) is probably synonymous, based on
examples in which the intermediary lirae are specially well developed and the
sutures abnormally sunken.
Bullia diluta (Krss.)
Fig. 27(d)
1848. Krauss. Stidafrik. Moll., p. 121 (Buccinum d.).
1874. Von Martens. Jahrb. D. Malak. Ges., i, p. 137, pl. 6, fig. 4 (var. mediolaevis).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 53, pl. 35, fig. 5 (aepynota, subturriform).
1915. id. ibid., p. 54, pl. 3, fig. 2 (alfredensis).
Probable other synonyms are Turton’s: albanyana (subturriform), rietensis,
zenobia, subventricosa (subturriform), spectrum, scitula.
Aperture 13-1? (half-grown) to 2 times in spire. Angle of spire 20°-25°.
Apical whorl (protoconch) diam. 0-75 mm. Whorls (total) 9. Profile of whole
shell straight. Parietal callus thin, extending up @-7 of length of preceding
whorl and visible from (5th) 6th whorl onwards (best seen in worn specimens!).
Spiral striae over whole whorl from 3rd whorl onwards, c. 15 on last whorl,
but often with fine intermediaries, striae closer together on upper part of
134 ANNALS OF THE SOUTH AFRICAN MUSEUM
whorl; 5-6 additional spaced striae on base. Columellar glaze thin, not
extensive, not covering rostrum. 29 X 10 mm.
Operculum and animal unknown.
Cream or fawn, with a series of orange-brown spots in upper third of
whorls, continued downwards as axial streaks (as if the spot of colour had
‘run’), the costa on base from columella to end of aperture orange-brown,
sometimes showing through at the suture of following whorl, the ground-colour
above the spots often paler (whitish) than the lower part of whorls.
Natal (Krauss); Port Elizabeth (Sowerby, Bartsch); Port Alfred (Bartsch,
Turton). Port Alfred, East London, Port St. Johns, Scottburgh (Natal),
Durban, Tongaat (30 miles N. of Durban) (S. Afr. Mus.). Delagoa Bay (U.W.).
False Bay (von Martens: var. mediolaevts).
Remarks. A common species with several minor variations in coloration
and convexity of the whorls, sometimes subturriform (aepynota, subventricosa,
scitula), which have been given names as distinct species.
The specimens described by von Martens as var. mediolaevis from the
Fritsch collection were stated to have come from False Bay. But there is no
other record from any locality west of Algoa Bay; perhaps Dr. Fritsch obtained
the specimen from a friend.
Bullia natalensis (Krss.)
Figs. 25(c¢), 27(h), 28
1848. Krauss. Stidafrik. Moll., p. 121, pl. 6, fig. 16 (Buccinum n.).
Aperture 14—14 in spire. Diameter of protoconch 0:5, of 1st whorl 1-25,
of and 1°5, of 3rd 1-75 mm. Whorls (total) 8. Profile straight, slightly concave
near apex. Parietal callus thin, visible but very narrow on 3rd whorl, on 4th
extending up $—} length of preceding whorl, on 5th 4, on 6th 4, and on 7th
and 8th %. First 3 whorls smooth; fine axial slightly curved plicae on 4th and
5th, somewhat irregular, becoming stronger on later whorls, where they form
a strong crenulation at the suture, but obsolescent on middle and lower part
of whorl, about 22 on last whorl. Spiral striae on 4th and following whorls,
about 13 on 4th whorl, 15 on 5th, closer together at top of whorl, on later
whorls less conspicuous and concealed by the increasing width of the callus,
3-4 additional inconspicuous striae on base. Columella evenly curved, without
keel or bend; glaze thin, covering about 4-4 base. 34 x 13 mm. (8 whorls).
A larger specimen 41 X 17 mm. (apex worn, 4 whorls remaining, probably
43-44. mm. long when perfect).
Operculum nearly filling aperture, ovate, apex incurved, margins smooth,
5 X 2°5 mm. in shell 15 mm. with 6 mm. aperture.
Cream or flesh-coloured, sometimes slightly livid in middle of whorl,
basal costa yellowish, aperture orange within, pale at margin of outer lip,
operculum amber.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 135
Foot with antero-lateral corners shortly pointed, posteriorly with 2 tails,
no eyes. Radula with 55 (14 mm. sheli)—65 rows, central plate slightly wider
than lateral plate, its base 3-34 times as wide as long, with 9-10 cusps; lateral
plate with inner cusp bifurcate, the outer prong smaller and more slender than
the inner (median) prong. Animal pale flesh-colour.
Fossil: raised beach, alt. 375 ft. Durban-Umgeni (Geol. Survey).
Dead: Durban (Krauss). Durban and Tongaat (S. Afr. Mus.).
Living: Durban (S. Afr. Mus. coll. K.H.B.); Umhlali and Karridene
(Natal) (U.C.T.), Delagoa Bay (U.W.). |
4) eel
ab iap
pf
Fic. 28.
Protoconch and upper whorls of (eft) Bullia natalensis (Krss.), and (right)
B. mozambicensis Smith. Length of portion figured 3 mm. and 2 mm. respectively.
Remarks. Distinguished from mozambicensis by the apical three smooth
whorls being more elongate and papilliform, and by the sculpture of the early
whorls being cancellate but not pustulose, and by the lack of spiral striae on
base of last whorl. The operculum very nearly fills the aperture, and relatively
is larger than in any other South African species.
The parietal callus band is also, relatively, wider than in other South
African species.
This species does not, according to the few records, occur south of Durban.
Aberr. gigant. In S. Afr. Mus. (ex coll. Ross-Frames) is one very large
shell said to have come from the mouth of the Limpopo River (Chai Chai,
Portuguese East Africa), together with one undoubted mozambicensis. It
measures 60 X 24 mm.; 6 whorls remain, the apex is worn and 2, possibly 3,
whorls are missing. It agrees in sculpture and shape of columella with natalensis.
The parietal callus extends only 4 in the 2nd remaining whorl, but rapidly
widens to 4 on the 3rd and equally rapidly to $ on the 4th, leaving only the
sutural crenulations exposed; near the end of the 4th whorl it drops abruptly
to % on 5th, and continues thus on the 6th.
136 ANNALS OF THE SOUTH AFRICAN MUSEUM
The sutural diameter of the 8th whorl of the 34 mm. shell is 10 mm.; if
the 41 mm. shell (recorded above) be assumed to have had 84 whorls the
diameter of the 8th whorl is also 10 mm.; and the diameter of the 84th of this
41 mm. shell is approximately the same (12 mm.) as that of the assumed 84th
whorl of the giant 60 mm. shell. The latter may therefore be reasonably
assumed to have had originally 9, possibly 94, whorls.
Until connecting sizes are found this ‘outsize’ shell may be regarded as an
example of gigantism.
Recently an even larger shell has come to hand, collected by Dr. L. Kent
(Geological Survey) somewhere between Port Shepstone and Durban. It
measures 65 <X 26 mm.; only 44 whorls remain, the 34 or 44 apical whorls
being missing. The callus on the last two whorls occupies one half of the
preceding whorl.
Bullia mozambicensis Smith
Figs. 25(d), 27(z), 28
1877. Smith. Proc. Zool. Soc. Lond., p. 719, pl. 75, fig. 18 (not good).
1894. Sowerby. 7. Conch., vii, p. 368 (pustulosa).
1897. id. Append. Mar. Sh. S. Afr., p. 5, pl. 6, fig. 1 (pustulosa).
1931. Tomlin. Ann. Natal Mus., vi, p. 430.
Aperture 14 (—14) in spire. Diameter of protoconch 0:5, of 1st whorl 0-75,
of end 1°5, and of 3rd 1-75 mm. Whorls (total) 9. Profile straight, slightly
concave near apex. Parietal callus thin, very narrow on 4th whorl, on 5th
extending up 4-4 length of preceding whorl, on 6th—oth whorls up to 4 or
nearly 4, never exceeding 4. First three whorls smooth; close-set regular
slightly curved axial plicae on 4th and following whorls, from suture to suture
on early whorls, but on (7th) 8th and goth becoming obsolete on middle and
lower part of whorl, forming a strong crenulation at the suture, c. 22 on 4th,
30-32 on 5th and later whorls, but becoming irregular and more or less
coalescent on 8th and goth whorls. Spiral striae 8-g on 4th and following
whorls, 9-10 on gth whorl, deep, cutting the axial plicae into conspicuous
pustules on 4th—7th whorls (not well shown in Smith’s figure), but on 8th the
pustules are flattened and the sculpture consequently only cancellate, middle
and lower part of 9th whorl with only spiral striae; 4-5 additional striae on
base; the pustules and striae on the lower part of the whorls not completely
concealed by the callus. Columella with a somewhat pronounced bend at the
anterior canal; glaze thin, covering not more than 4 of base. 43 x 18 mm.
Operculum not filling aperture, ovate, apex incurved, margins smooth,
6-5 X 3 mm. in 31 mm. shell with 13 mm. aperture.
Cream or flesh-coloured, sometimes slightly livid on middle of whorls,
aperture orange within but pale at margin of outer lip, operculum amber.
Foot with antero-lateral corners shortly pointed, posteriorly with 2 tails,
no eyes. Radula with 80 rows, central plate 14-14 times as wide as lateral
plate, its base about 4 times as wide as long, with 11-12 (13) cusps, lateral
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 137
plate with inner cusp bifurcate, outer prong more slender than inner prong.
Animal pale greyish-buff.
Dead: Quelimane (Smith); Durban (Sowerby); Durban and Tongaat
(S. Afr. Mus.); mouth of Limpopo Ruver (Chai Chai, Portuguese East Africa)
(S. Afr. Mus. coll. Ross-Frames).
Living: Chinde, mouth of Zambesi River (S. Afr. Mus. coll. K.H.B.).
Remarks. ‘The differences between this species and natalensis have been
given above. The length of the first three whorls is here less, and the shape
therefore more mamilliform.
Bullia digitalis Meuschen
Figs. 25(¢), 27(7 ), 29
1787. Meuschen. Mus. Gevers., p. 296.
1816. Lamarck. Tabl. Encycl. Meth., pl. 400, figs. 4 a, b, and Liste, p. 2 (Buccinum achatinum,
nom. et fig.).
1846. Reeve. Conch. Icon., ii, Bullia sp. 4, pl. 4 (digitale), sp. 14 (sulcata), sp. 17 (semiflammea).
1847. id. ibid., sp. 22 (semiusta).
1885. Euthyme. Bull. Soc. malac. Fr., ii, p. 237 (capensis).
1903. Von Martens. D. Tiefsee Exp., vii, p. 53 (digitata [sic] et alia).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 54, pl. 35, fig. 4 (almo).
1921. Sowerby. Proc. Mal. Soc., XIv, p. 125, text-fig. (dulcis).
1922. Tomlin. 7. Conch., xvi, p. 260.
1923. Odhner. Géteb. K. Vet. Handl., xxvi, 7, p. 6.
1932. Turton. Mar. Sh. Pt. Alfred, pp. 65, 66, pl. 15 (digitalis and vars.).
1932. id. ibid., p. 65, pl. 15, no. 470 (soluta Gmelin, subscalariform).
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 22.
1937. Peile. loc. cit., p. 184 (radula) (achatina).
Aperture 14-12 (ocasionally 2) times in spire. Angle of spire 30°-35°.
Apical whorl (protoconch) diam. 0-6 mm. Whorls (total) 10. Profile of whole
shell straight. Parietal callus thin, visible from 4th whorl, extending up about
@ length of preceding whorl. Spiral striae from 3rd whorl onwards over whole
whorl, but often indistinct and obsolescent on middle and lower part of whorls,
very fine, close together, minutely crinkly due to intersection with the fine
growth-lines; 5-6 additional stronger and widely spaced striae on base.
Columellar glaze thin, extending over about half base. 52 x 21 mm., smallest
examined 4 X 1°75 mm.
Operculum triangular, both margins strongly serrate, but often worn
nearly smooth, apex narrow, concave between two little points, 6 x 3°75 mm.
in 45 mm. shell with 16 mm. aperture.
Variously coloured: uniform cream, fawn, brown (achatina), sometimes
with livery tinge (sulcata), plumbeous or violaceous; upper half of whorls cream,
lower half brown (semiusta); cream with a peripheral series of orange-brown
spots which extend downwards (as if the colour had ‘run’) as axial streaks and
flames (semi-flammea, dulcis), sometimes the spots are absent and the axial streaks
may be separate and distinct, or they may be coalescent into a more or less
solid band, thus approaching the semiusta pattern; the apex is usually white,
138 ANNALS OF THE SOUTH AFRICAN MUSEUM
even in achatina; sometimes the early whorls may be
brown with the later ones becoming gradually uniform
cream. Operculum amber.
Foot with antero-lateral corners acute, posteriorly
with 2 tails, no eyes. Radula with 75-85 rows, central
plate about 14 times wider than lateral plate, its base
nearly 4 times as wide as long, with 15-17 cusps, and
often a minute denticle externally on one or both sides,
lateral plate with inner cusp bifurcate, outer prong
smaller than the inner prong.
Animal pale flesh-coloured, foot sometimes plum-
beous or violaceous.
Fossil, Quaternary: Angra Junta, South West Africa
(Haughton).
Dead: Angra Pequena (Liideritzbucht), Olifants
River mouth, False Bay, Port Elizabeth, Port Alfred
(auct). Table Bay, Plettenberg Bay (S. Afr. Mus.). Off
Cape Morgan, 36 fathoms (S. Afr. Mus. P.F. coll.).
Living: False Bay, littoral and 9 fathoms, Port Eliza-
Fic. 29. beth, littoral (S. Afr. Mus.); Hydra Bay (Danger Point) |
Bullia digitalis Meus- (Odhner, presumably living); Saldanha Bay and Lam-
chen. 14mm. specimen ?
fom Stil Bayito ahow bert’s Bay, 15 metres (U:@.1:);
abnormal development Remarks. ‘The earlier records from Liideritzbucht
haat a alee might have been open to doubt, but have been con-
like shoulder. Callus firmed by Tomlin (1922). The Olifants River, Lambert’s
SHPO Bay and Saldanha Bay records connect with Table Bay,
but there is a considerable gap between the first mentioned and Liuderitz-
bucht. The species has not been reported from Port Nolloth. At Lambert’s
Bay U.C.T. found it at 15 metres depth, but not on shore.
Examples with the semiflammea-dulcis colour pattern are liable to be con-
fused with diluta, but may be distinguished by the shorter spire relatively to
the aperture, and the fine spiral striae.
Subscalariform aberr. (fig. 29). A 14 mm. specimen (Still Bay, coll. Muir)
has the first 5 whorls normal, but the following two with well-marked shoulders.
Below the shoulder the profile of both whorls is straight, and thus the width of
the last whorl (5-5 mm.) is scarcely greater than that of an equal-sized normal
example. The 4th and 5th whorls show the peripheral series of spots, but the
last 2 whorls are uniform cream.
The change to shouldered whorls was caused by an injury. Instead of the
parietal callus being plastered as a thin layer against the preceding whorl, only
its edge adjoins the preceding whorl (forming a ‘false’ suture), the rest of the
layer projects outwards forming the shoulder. A later, less severe injury has
resulted only in an irregular joint, without further upsetting the animal’s
organization.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 139
Bullia rhodostoma (Gray MS.) Rve.
Figs. 25(g), 27(*)
1847. Reeve. Conch. Icon., iii, Bullia no. 25.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 55 (polita ? non Lam.).
1937. Peile. loc. cit., p. 184 (radula).
Aperture 14-1} times longer than spire (juv.), subequal to spire (half-
grown), 17 in spire (fully grown). Angle of spire c. 40°. Apical whorl (proto-
conch) diam. 0:75 mm. Whorls (total) 9. Parietal callus thin, visible from grd
whorl, extending up 4, sometimes almost 4 length of preceding whorl. Spiral
striae from 3rd whorl onwards, over whole whorl on 3rd and 4th but on later
whorls confined to upper part, obsolescent on middle and lower parts, even on
the upper part rarely more than half-a-dozen very fine striae, often indistinct
or obsolete, no additional striae or only very fine and indistinct ones on base
(see Remarks) ; growth-lines may produce a fine minute crinkling at the sutures.
Columellar glaze thin, extending at most only halfway across base. 47 x
‘Ig mm., smallest examined 4 xX 2°75 mm.
Operculum ovate, apex incurved, margins smooth, 9 X 3°75 mm. in 39
and 41 mm. shells with apertures 17-18 mm.
Cream or flesh-coloured, lower part of whorls sometimes darker, some-
times paler, due to the parietal callus, costa on base orange, aperture deep
orange within but paler at margin of outer lip, varices (if present) yellow or
orange; operculum amber.
Foot with antero-lateral corners shortly pointed, posteriorly with 2 tails,
no eyes. Radula with 55-60 rows (40-45 in 17 mm. shells), central plate only
a little wider than lateral plate, its base about 3 times as wide as long, with
II—13 cusps, often a minute denticle externally on one side or the other, lateral
plate usually with 2 smaller cusps on the outer margin of its inner cusp, on one
side sometimes with only one, sometimes with 3 (cf. Peile).
Animal pale greyish or flesh-coloured.
Dead: Port Elizabeth, Port Alfred (Sowerby, Bartsch, Turton). Pringle
Bay (east side of False Bay), and Still Bay (S. Afr. Mus.).
Living: False Bay, littoral; Mossel Bay; Port Elizabeth, littoral; Durban,
littoral (S. Afr. Mus.). Delagoa Bay (U.W.).
Remarks. ‘The Durban specimens conform with the above description in
all respects, including the animal with its radula, but the spiral striae are more
distinct than in the majority of examples from farther west; in one specimen
29 mm. long there are about 20 striae on the upper part of the last (8th) whorl
and 8-10 at bottom of base; the largest specimen 33 mm. also with 8 whorls,
has fewer striae above but more below, and faint traces of striae in the middle
of the whorl.
I have seen only one juvenile from Delagoa Bay, but the radula agrees.
In the Pringle Bay specimens the outer lip at irregular intervals has been
thickened to form a varix; in one specimen one varix on penultimate whorl
140 ANNALS OF THE SOUTH AFRICAN MUSEUM
and about 10 on last whorl, the latest-formed 4-5 close together; in another
specimen one varix on penultimate whorl and 4 on last; three others have 1-4
varices on the last whorl only; one other specimen has only one feeble varix
near margin of outer lip. Specimens from other localities very rarely produce
varices.
The specimen which Bartsch referred to ‘folita Lam.’ is probably a
rhodostoma. (The West African folita Lam. = muiran Brug. is a Dorsanum, see
Dautzenberg 1910.)
Bullia laevissima (Gmelin)
Figs. 25(h), 27(/)
1816. Lamarck. Tabl. Encycl. Meth., pl. 400, figs. 1 a, b, and Liste, p. 2 (Buccinum laevigatum,
nom. et fig.).
1886. Watson. Challenger Rep., xv, p. 190 (laevigata) (references).
1903. Von Martens D. Tiefsee Exp., vii, pp. 29, 53 (laevigata).
1922. Tomlin. 7. Conch., xvi, p. 260.
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), pp. 22, 46, pl. 5, fig. 3.
19392. ‘Gevyers. (ibid. p. 74.
1932. Turton. Mar. Sh. Pt. Alfred, p. 66, pl. 16, no. 482 (juv.), and var. globulosa.
1937. Peile. loc. cit., p. 184, fig. 19 (radula) (Leiodomus I.).
Aperture at first subequal to spire, but at an early stage becoming longer
than spire, in adult spire 14 in aperture. Apical whorl (protoconch) diam.
0-75 mm. Whorls (total) 7, perhaps 8, broadening rapidly from grd or 4th
whorl onwards. Parietal callus thick, intervening between suture and preceding
whorl, and extending up about 4 length on 4th whorl (sometimes beginning to
show on 3rd), increasing to 4, often to 2 or even 2 length of preceding whorl in
large shells. On 2nd whorl a spiral stria at top of whorl, becoming a sulcus on
3rd, rather broad but shallow on 4th and 5th whorls, but narrowing and
obsolescent on 6th, on 7th only a slighi indentation below the rounded shoulder;
2-3 (sometimes 4) striae on lower part of base in juveniles, but in older examples
only the lowermost one persisting, and that one obsolete in very large shells
(over 35 mm. long). Columellar glaze thick and extending over whole base.
55 X 34 mm., another with worn apex 57 mm. long; Tryon gives 75 mm.;
smallest specimen examined 4°25 X 2°75 mm.
Operculum very small, subtriangular (similar to that of annulata but much
smaller), or narrow oval or cuneiform, apex truncate, margins smooth,
5 X 4 mm. or 5 X 2 mm. in 42 mm. shells with 25 mm. apertures, and
4°5 X 2°5 mm. in 55 mm. shell with 35 mm. aperture.
Cream, fawn, or greyish, with 2-3 faint spiral darker bands, the band
near the suture more visible than the other(s); columella suffused with pink
or madder-brown; operculum amber-brown. |
Foot with antero-lateral corners very shortly pointed, posteriorly with 2
small tails, no eyes. Radula with c. 80 rows, central plate 14-14 times as wide
as lateral plate, its base 4-44 times as wide as long, with (18) 19-22 cusps,
often a minute denticle externally, lateral plate with inner cusp bifurcate,
y
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA I41
symmetrical, or the outer prong bifurcate on one side, or bifurcate on one side
and trifurcate on the other (cf. Peile).
Animal purplish-brown or livery-pink, edge of foot pale.
Fossil, Quaternary: Bogenfels and Angra Junta, South West Africa
(Haughton) ; Sedgefield near Knysna (Martin); Recent: Cape Cross salt-pan,
South West Africa (Gevers).
Dead: Liideritzbucht, Olifants River mouth, Table Bay, False Bay, Algoa
Bay, Port Alfred (auct.).
Living: Simon’s Bay (False Bay), 15-20 fathoms (Watson). Saldanha
Bay, 10-14 fathoms; Table Bay; False Bay, 10-22 fathoms, Mossel Bay and
off Cape St. Blaize, 17-19 fathoms (S. Afr. Mus. P.F. coll.).. Lambert’s Bay,
15 metres; Saldanha Bay, False Bay and Algoa Bay (U.C.T ); 34° S 25° 46’ E.,
41 fathoms (U.C.T.).
Remarks. Smith (1903) said laevigata Chemn. was not published by a
binomial writer, but Watson (1886) quotes “Buccinum laevigatum’? Chemn. 1780,
with full reference, and laevissima Gmelin 1790.
This is the only South African species in which the aperture exceeds the
spire in length (except in very juvenile).
Juveniles less than 10 mm. in length are not always distinguishable with
certainty from those of annulata, but are usually proportionately broader and
the length of the aperture exceeds that of spire.
Bullia callosa (Gray) Rve.
Fig. 27(m)
1828. Gray in Wood. Suppl. Ind. Test., p. 12, pl. 4, fig. 14 (Buccinum c.) (fide Dautzenberg,
1912. Sherborn, Ind. Anim., says nom. et fig.).
? 1839. Gray. Kool. of the ‘Blossom’, p. 127 (semiplicata).
1847. Reeve. Conch. Icon., tii, Bullia sp. 24.
1884. Fischer. Man. Conchyl., p. 636, pl. 5, fig. 4 (Buccinanops semiplicata Gray).
1889. Sowerby. 7. Conch., vi, p. 6 (var. sulcata).
1903. Von Martens. D. Tiefsee Exp., vii, p. 53.
1912. Dautzenberg. Ann. Inst. ocean., vol. v, fasc. 3, p. 34 (Dorsanum c.).
1932. Gevers. Tr. Geol. Soc. S. Afr., xxxiv (1931), p- 74.
an
Aperture subequal to spire. Apical whorl (protoconch) diam, 0-75 mm.
Whorls (total) 7, turreted (pagoda-like), profile straight, at least on later
whorls. Parietal callus thick, often very thick, intervening between suture and
preceding whorl, and extending up about 4 length, when strongly developed
extending not farther upwards but outwards to form a rounded shoulder above
the suture of following whorl. On 3rd whorl a spiral stria at top of whorl (some-
times traceable on later part of 2nd whorl) becoming on 4th and later whorls
a sulcus followed by a stria, sometimes 2-3 additional striae traceable but
rarely distinct, 4-5 additional ones on base. Columellar glaze thick and
extending over whole base. 37 X 19 mm. (worn); Turton: 45 mm. long.
Smallest specimen examined 3 X 1°75 mm.
142 ANNALS OF THE SOUTH AFRICAN MUSEUM
Operculum small, narrow triangularly ovate, margins smooth, 5°5 x
2°5 mm. in 35 mm. shell with 20 mm. aperture.
Brown or purplish-brown, the callus band darker, openciites amber.
Beach specimens cream, fawn or pale brown, the callus forming a brown spiral
band and a brown patch on base. !
Foot with antero-lateral corners shortly pointed, posteriorly with 2 small
tails, no eyes. Radula with c. 90 rows, central plate 14 times as wide as lateral
plate, its base about 24 times as wide as long, with 13-15 cusps; lateral plate
with inner cusp bifurcate, outer prong more slender than the inner (only one
radula available).
Animal purplish-brown, sole darker than upper part of foot.
Fossil, Recent: Cape Cross salt-pan, South West Africa (Gevers).
Dead: Liideritzbucht, Port Elizabeth, Port Alfred (von Martens, Sowerby,
Turton). Still Bay (S. Afr. Mus. coll. Muir); Durban and Tongaat (30 miles
N. of Durban) (S. Afr. Mus.).
Living: Mossel Bay and off Cape St. Blaize, 17 fathoms (S. Afr. Mus.
Pat icoll)). Algoa Bay (UU. G=0)).
Distribution. Mossamedes, 15-20 metres (Dautzenberg).
Remarks. Juveniles 7-8 mm. long (5 whorls) usually, and 5 mm. long
(4 whorls) sometimes, are recognizable by the brown callus band; smaller
examples in which the callus is as pale as rest of shell are very similar to
juveniles of annulata but narrower.
The Durban and Tongaat examples—up to 38 mm. long—show the sub-
sutural sulcus and 3-4 striae very clearly (var. sulcata), except on the last (7th)
whorl of the largest specimen
On the south coast this species has not been recorded from farther west
than Still Bay, not even from False Bay. There is therefore a big gap in the
distribution until one comes to Liideritzbucht and Cape Cross on the west
coast, and farther north Mossamedes.
Presumably the south and west coast examples are conspecific. Live
specimens from the west coast would be useful for comparison, because the
South African callosa is a true Bullia, i.e. without eyes and with two tails, not a
Dorsanum as defined by Thiele (1929).
Gen. Adinopsis Odhner
1923. Odhner. Géteb. K. Vet. Handl., xxvi, 7, p. 15.
1929. Thiele. Handbuch, i, p. 740 (addenda).
1937. Peile. Proc. Mal. Soc., xxii, p. 186 (radula).
Shell in general resembling Bullia, profile slightly concave, apex blunt,
no parietal callus, columellar glaze narrow, outer lip plicate within.
Operculum apex incurved, margins smooth.
Animal with eyes, foot with 2 tails posteriorly. Radula lateral plate with
2 cusps, the inner one not bifurcate.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 143
Adinopsis skoogt Odhner
1923. Odhner. loc. cit., p. 15, pl. 1, figs. 11-14.
1937. Peile. loc. cit., p. 186, fig. 24 (radula).
Length 25 mm. Brown with two pale spiral bands.
Port Alexander, Angola, 16 fathoms.
Fam. VOLEMIDAE
1929. Thiele. Handbuch, 1, p. 319 (Galeodidae).
1952. Bayer. ool. Med., xxxi, no. 25, p. 265.
If Volema Bolten-R6ding 1798 is admitted as a valid genus, the family
name should be Volemidae. Bayer pointed out that Galeodes Bolten-R6éding is
preoccupied, and therefore adopted Melongena. He included as a section of
this genus the earlier Volema; it would seem more correct to make Volema the
genus, with sections Volema s.s. and Melongena.
Gen. VoLeMA Bolten-Réding
1798. Bolten-Roding. Mus. Bolten. (2), p. 57.
1817. Schumacher. Essai. . . vers testacés, pp. 64, 212.
1929. Thiele. loc. cit., p. 320 (Galeodes, non Olivier, 1791).
1952. Bayer. loc. cit., p. 265 (Malongena).
Volema paradisaica (Martini-Reeve)
Fig. 30(a), (5)
1777. Martini. Syst. Conch. Cab., iii, p. 202, figs. (full reference in Bayer) (Pyrum p.).
1790. Gmelin. Linn. Syst. Nat., ed. 13, p. 3484, no. 56 (Buccinum pyrum).
1847. Reeve. Conch. Icon., iv, Pyrula, no. 17.
1933. Krige. Tr. Geol. Soc. S. Afr., xxxv (1932), p. 52.
1952. Bayer. loc. cit., p. 276 (pirum, [sic]).
1952. Braga. Anais F. Invest. Ultramar., vii, 3, p. 73, pl. 2, fig. 4 (Melongena p.).
Protoconch 2 whorls, diam. 1-3, alt. 1:5 mm., smooth, with a few feeble
plicae before the not sharply defined junction with Ist postnatal whorl; the
latter with 10-11 peripheral knobs.
Radula with c. 120 rows, central plate longer than wide, anteriorly con-
cave, posteriorly convex, outer cusp as long as basal plate, middle cusp much
smaller; lateral plate as wide as length of basal plate of central, outer cusp
twice as long.
Fossil, raised beach, alt. 375 ft. at Durban-Umgeni (Geol. Survey).
Durban Bay, Delagoa Bay, Mozambique (auct. wide Bayer).
Durban Bay, one with periostracum (S. Afr. Mus.); Inhambane, juv.
EOLe. 1).
Living: Delagoa Bay (S. Afr. Mus. coll. K.H.B. and U.W.).
Distribution. Red Sea, Zanzibar, Mombasa, Ceylon, East Indies.
144 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. Bayer gave references to early descriptions.
Sowerby’s record from Port Elizabeth can scarcely be based on even a
beach specimen, certainly not a living one; it was transported more probably
by a collector than by a marine current.
Am) <0
Fic. 30.
(a), (b) Volema paradisaica (Mart. Rve.). Protoconch; central and lateral radula plates.
(c) Engina mendicaria (Linn.) central and lateral radula plates.
Fam. BUCCINIDAE
Tomlin (1932) described two species in the genus Glypteuthria solely on
conchological characters. One of these (capensis) has since been shown by the
radula to belong to Fusivoluta, and the other (solidissima) conforms in all essentials
with Afrocominella elongata. This Antarctic genus, therefore, has no place in the
South African fauna-list.
Tomlin (1932) also proposed a new genus Charitodoron in this family. One
of Tomlin’s species is, in my opinion, synonymous with Columbella agulhasensis
Thiele, which raises the possibility that the genus may be Columbellid, or even
Mitrid (see pp. 146 and 51,52).
The P.F. obtained no examples of the genera Phos. Metula, or Pisania, and
these genera are not discussed here.
A most unlikely species to be found on the Natal coast or off the Cape is
the New Zealand Szphonalia mandarinus (Duclos), recorded by Krauss (1848)
and Studer (1889). The Gazelle specimens from off the Cape were probably
Fusiwoluta (see p. 30); and Krauss’s specimens probably a Fasciolaria (see
P: 77): |
‘Cominella’ species. The following species are transferred to Muricid
genera: aculispira Sow., fuscopicta Turton, punciurata Sow., unifasciata Sow.,
wahlbergi Krss.; sulcata Sow. has been transferred to Daphnella, and angusta Sow.
might well be compared with Mangilia ponsonbyi ( Turritidae).
Gen. CHARITODORON Tomlin
1932. Tomlin. Ann. S. Afr. Mus., xxx, p. 164.
1943. id. 7. Conch., xxii, p. 50.
Shell fusiform, thin-walled, aperture shorter than spire, whorls with spiral
grooves or striae, axial ribs present on early whorls, sometimes feeble, or
9
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 145
entirely absent; no parietal callus, columella straight, canal short and broad,
outer lip thin, internally smooth; periostracum thin. Operculum and animal
unknown.
Remarks. This genus has been found not only off the Cape, but also in
deep water off East London; its occurrence in deep water off the Natal coast,
therefore, would not be very surprising. In fact one shell, at first thought to be
a Charitodoron, was recently found while searching through the P.F. bottom-
samples.
This 4-whorled shell would have been referred to C. pasithea in spite of the
remoteness of Natal from the type locality of this species, because of its close
similarity in measurements with the upper 4 whorls in Tomlin’s figure (which
is X 2) and its agreement with his description. Fortunately, however, the shell
contained the animal, .and the radula showed it to be a Mitrid (see supra,
Dibaphus, p. 51). :
The remoteness of the two localities thus seems, conversely, to override
the conchological similarities, and to be a positive reason for not identifying
the Natal shell with C. pasithea. At least, until a radula is obtained from an
undoubted species of Charitodoron, it is safer to leave this genus where Tomlin
placed it. Quite possibly it may eventually be removed to the Columbellidae in
which Thiele described his agulhasensis (syn. aglaia Tomlin, see p. 146).
; Key to species
1. Early whorls cancellate.
meeater whorls punctate-siriate .9 5 5 6. 6 6 eee ees 8) ow pe Cuphrosyne
Pee ae raw Ors SIMOOLME he ts.) | she, a Gt ye, lps vos aa: «) egulhasensis
c. With deep spiral grooves, cancellate sculpture on last whorl feeble . pasithea
Pee MvHons witospiral striae only << 5 . 2. ws ee ew el Ue. Bhalia
Charitodoron euphrosyne ‘Tomlin
1932, Vomlin. loc. cit., p. 167, fig. 8.
Protoconch 24 whorls, diam. 1, alt. 1:25 mm., smooth. Postnatal whorls
6; (1st corroded) 2nd and 3rd cancellate with axial ribs and spiral lirae, the
former gradually petering out on 4th whorl, 5th and 6th whorls with punctate
striae only, on 2nd and grd whorls 6-7 striae, on 4th 7-9, on 6th 8-10, on
base 10-12 (excl. those on rostrum), additional stronger striae becoming deeper
and broader grooves. 30 (protoconch missing) xX I0 mm.
White with pale yellowish periostracum.
S.W. of Cape Point, 660-700 fathoms (Tomlin); off west coast of Cape
Peninsula and south-west of Cape Point, 130-210 fathoms, all dead (S. Afr.
Mus. P.F. coll.).
Type in 8S. Afr. Museum.
Remarks. Only one specimen has a complete protoconch; and in all the
specimens corrosion has affected the 1st, and often the 2nd and partly the 3rd
whorl. The protoconch appears to have been smooth.
146 ANNALS OF THE SOUTH AFRICAN MUSEUM
Charitodoron agulhasensis (Thiele)
1925. Thiele. D. Tiefsee Exp., xvii, p. 173, pl. 30 (18), fig. 20 (Columbella a.).
1932. Tomlin. loc. cit., p. 169, fig. 9 (aglaia).
Protoconch 24 whorls, diam. 1 mm., smooth. Postnatal whorls 7 (6 in
Type of aglaia); 1st and 2nd whorls cancellate with numerous axial ribs and
6 spiral striae; 3rd (and sometimes part of 4th) obscurely punctate-striate;
later whorls smooth except for the fine slightly arcuate growth-lines, in some
lights 4-6 extremely faint spiral lirae may be visible; base with about 12 (excl.
those on rostrum) additional grooves. Type (aglaia): 26 X 9 mm., 31 X 10 mm.
Type (agulhasensis): 17 X 5 mm.
| White with pale yellowish periostracum, with indications here and there
of slightly darker axial flames.
35° 16'S. 22°26 E., 155 metres (Thiele); exact locality ay) Menaam:
Cape St. Blaize N. x E. distant 73 miles, 125 fathoms; 36° 40’ S. 21° 26 E.,
200 fathoms; off Cove Rock (East London area), 80-130 fathoms; all dead
(S. Afr. Mus. P.F. coll.).
Type of aglaia in 8. Afr. Museum.
Remarks. Toralin’s description says there are 9 whorls, but the Type has
only 8 (total), the tip of the protoconch being worn away.
There are three smaller specimens in 8. Afr. Mus. from the same haul as
the Type. As in euphrosyne the apices are corroded.
The Cape St. Blaize specimen is larger than the Type, but is badly cor-
roded, only the apex and body whorl and connecting columella remaining.
Protoconch 24 whorls plus 7 whorls. Periostracum greyish-brown. Six smaller
specimens were taken in the same haul.
The Cove Rock specimen, entered in the Museum Register, seems to be
missing from the collection.
It is a pity to have to displace Tomlin’s specific name, but the synonymy
seems clear; a 19 mm. specimen in S. Afr. Mus. corresponds exactly with
Thiele’s description and figure.
Charitodoron pasithea ‘Tomlin
1943. ‘Tomlin. loc. cit., p. 50, fig.
Protoconch eroded, about 7 postnatal whorls; deep regular spiral grooves,
8 on penultimate whorl, slight arcuate axial ribs on upper whorls, hardly
traceable on body whorl. 21 x 8 mm.
Off Cape Point, 430-630 fathoms (Tomlin); same locality, down to 800
fathoms, all dead (S. Afr. Mus. P.F. coll.).
Remarks. ‘Tomlin said all the specimens were in one way or another broken
and mended; the figure shows a varix where the shell has been mended.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA LAY)
The above description is abstracted from Tomlin. None of the four
registered specimens (Tomlin said: ‘several’) are in S. Afr. Museum; they are
either still in the Tomlin collection, or were lost in transit during the war.
Charitodoron thalia Tomlin
1932. Tomlin. loc. cit., p. 169, fig. 10.
Protoconch 24 whorls, diam. 1:3 mm., smooth (but worn). Postnatal
whorls 5; whorls slightly shouldered (more so in the second specimen than in
the Type) immediately below the impressed suture; fine growth-lines, spiral
(non-punctate) striae on all whorls, 9 on early whorls increasing to 10-12 on
body whorl, c. 16 (c. 20 incl. those on rostrum) additional stronger grooves on
Mase 21 x 8mm.
White with pale cream periostracum.
Off Cape Point, 131 fathoms (Tomlin) ; off Gape Point, 800-900 fathoms;
off Buffalo River (East London area), 310 fathoms; all dead (S. Afr. Mus.
PF .’coll.).
Type in 8. Afr. Museum.
Remarks. A relatively broader species than either euphrosyne or agulhasensis.
Although the tip of the protoconch is worn smooth in both examples, I
find only 5 postnatal whorls. The protoconch is obviously larger than in
euphrosyne and agulhasensts.
The smaller specimen (not seen by Tomlin) has a small but distinct square
shoulder below the suture.
The East London specimen, entered in the Museum Register, seems to be
missing from the collection.
Gen. BABYLONIA Schl.
1822. Lamarck (Eburna, non Lamarck, 1801).
1838. Schliiter. Kurzg. syst. Verz. Conch., p. 18.
1929. Thiele. Handbuch, i, p. 312 (fig. 344 radula).
1929. id. ibid., p. 332 (<emiropsis, Olividae).
1937. Peile. Proc. Mal. Soc., xxii, p. 182 (systematic position).
1953. Kubo & Kondo. 7. Tokyo Univ. Fish., xxxix, p. 199, fig. 2 B (age determination based
on operculum).
1957. Yoshihara. ibid., xliii, p. 207, pl. 7, figs, C, D (spawning and egg-capsule).
Remarks. ‘Thiele in making a new genus (in the Olividae!) for the South
African species, evidently had not seen or had overlooked Sowerby’s 1902
record and figure. Even without recourse to the radula the species is obviously
a Babylonia, to which genus Peile restored it.
Babylonia papillaris (Sow.)
Fig. 31(a)
1825. Sowerby. Cat. Tankerville Coll., Append., p. xxii (Eburna p.).
1833. id. Conch. Illustr., Pt. 20, Eburna, no. 9, fig. (Eburna p.).
148 ANNALS OF THE SOUTH AFRICAN MUSEUM
1902. id. (3rd). Mar. Invest. S. Afr., ii, p. 93, pl. 2, fig. 3 (Eburna p.) (shell and animal,
operculum badly drawn).
1929. Thiele. loc. cit., p. 332 (<emiropsis p.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 31 (Eburna p.) and p. 31, pl. 6, no. 229 (millepunctata).
1937. Peile: loc. cit’, p..1é2, fie.15 (radula).
Protoconch 14 whorls, diam. at apex 1-5-2 mm., increasing to 2°5-3 mm.
Postnatal whorls 5. 47 (to end of columella) 50 (to base of aperture) <X 27 mm.
Smallest specimen seen 10 mm. long.
Operculum elliptical, concave, nucleus apical, 16 X 7mm. in 38 mm. shell.
White with reddish or orange-brown spots, varying from moderately
numerous (1-2 mm. diam.) to very numerous (0-5 mm. or less) (millepunctata) ;
sometimes larger spots near the suture and around base, and forming a central
band, small spots on rest of whorl; the large spots near suture usually larger
than others and oblique, flame-like; operculum yellowish-amber. Periostracum
yellowish.
Animal (as preserved) flesh-coloured; possibly orange when alive.
Wr) RLIN cae
Sl STy
Fic. 31.
Central and lateral radula plates of (a) Babylonia papillaris (Sow.); (6) Nassaria gracilis Sow.,
central plates from front, middle and hind portions of radula; (c) the same, from another
animal; (d) central plate of a Delagoa Bay specimen; (e) Afrocominella capensis (Dnkr.);
(f) A. elongata (Dnkr.); (g) Burnupena cincta (Bolten); (h) B. lagenaria (Lam.), central plate
only; (2), (7) Euthria queketti Smith, with malformation of the 77th (from the front) central plate.
Radula with 30 rows (Peile: 36 rows), central and lateral plates as in
Thiele’s and Peile’s figures, middle cusp on central plate a little longer than the
others.
Living: Algoa Bay, 24 fathoms (Sowerby, 1902). St. Sebastian Bay,
27 fathoms; Algoa Bay, 10-36 fathoms; off East London, 22-52 fathoms; off
Port Shepstone (Natal), 36 fathoms (S. Afr. Mus. P.F. coll.). False Bay
(U.C.T.). 34° 01'S. 25° 45’ E., 25 fathoms (millepunctata) (U.C.T.).
Dead: Natal (Smith); Port Alfred (Bartsch, Turton).
jack:
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 149
Remarks. The type of this species was in the Tankerville collection (now ?);
when describing it Sowerby recorded a second specimen then in Broderip’s
possession (now Brit. Mus.), formerly in the ‘African Museum’ (Bullock’s).
Mr. Galbraith of the British Museum informs me that there are two specimens
in the B.M., one of which may be a syntype.
Gen. ENcina Gray
1839. Gray. ool. of the ‘Blossom’, p. 112.
1840. Swainson. Treat. Malac., clii, p. 313 (Pusiostoma part).
1939. Peile. Proc. Mal. Soc., xxiii, p. 271 (radula).
Engina perlata (Kister)
1858. Kiister. Conch. Cab., p. 61, pl. 12, figs. 5, 6.
1895. Melvill. Proc. Mal. Soc., i, p. 226, pl. 14, fig. 12 (natalensis).
Radula as in mendicaria, but the external denticles on the central plate
are minute or obsolete; ¢. 98 rows. |
Living: Durban and Umhlanga (U.C.T.).
Dead: Natal (Kiister); Durban (Melvill); Tongaat and Scottburgh
(Natal), and Port St. Johns (S. Afr. Mus.).
Engina (Pusiostoma) mendicaria (Linn.)
Fig. 30(c)
1758. Linne. Syst. Nat., ed. 10, p. 731 (Voluta m.).
1859. Chenu. Man. Conchyl., i, fig. 1106 (Columbella m.).
1903. Von Martens. D. Tiefsee Exp., vii, p. 97 (references).
1911. Schepman. Siboga Exp. monogr., xlix, p. 3009, pl. 24, fig. 3 (radula).
Radula with ¢. 100 rows, central plate with 3 subequal major cusps, and
a small cusp externally on each side, lateral plate with outer cusp considerably
larger than the inner.
Living: Kosi Bay (U.C.T.); Delagoa Bay (U.W.); Mozambique Island
(S. Afr. Mus. coll. K.H.B., and U.W.).
Distribution. Mauritius, Seychelles, Madagascar; Indo-Pacific.
Engina marmorata (Rve.)
1846. Reeve. Conch. Icon., pl. 12, fig. 95 (Buccinum m.).
1848. Krauss. Siidafrik. Moll., p. 120 (Buccinum m.).
1880. Von Martens. Mauritius G Seychellen, p. 239 (Pisania m.).
1939. Peile. Proc. Mal. Soc., xxiii, p. 271, fig. 39 (radula).
Protoconch ? Postnatal whorls 7; axial ribs to on Ist whorl, increasing
to 18 on penultimate whorl and becoming feeble and obsolete on later part of
body whorl; crossed by spiral lirae 3 on Ist whorl, increasing to 8-9 on last
whorl, with one very fine intermediary between some of the pairs, ¢c. 12
additional lirae on base with intermediaries. Parietal callus dentiform,
150 ANNALS OF THE SOUTH AFRICAN MUSEUM
columella anteriorly keeled, outer lip internally plicate. 23 (protoconch
missing) X IO mm.
White, variegated with indefinite yellowish and brown patches.
Radula with up to 114 rows, central plate with 3 major and 2 minor
cusps, lateral plate with 2 cusps (Peile).
Natal (Krauss, and 8. Afr. Mus.); Port Elizabeth and Natal (Sowerby).
Distribution. Mauritius, Seychelles, Philippine Islands.
Remarks. Peile shows that this species cannot be included in Pisania or
Cantharus on account of its radula, which resembles that of Engina. He transfers
it to Engina with a query.
Gen. CANTHARUs Bolten-Réding
1798. Bolten-Réding. Mus. Bolten (2), p. 132.
1834. Gray in Sowerby. Gen. Sh., no. 42 (Pollia).
1840. Swainson. Treat. Malac., Ixxiv, p. 302 (Tritonidea).
Cantharus carinifera (Kister)
1858. Kiister. Conch. Cab., p. 63, pl. 12, figs. 9, 10.
1901. Smith. 7. Conch., x, p. 111, pl. 1, fig. 23 (natalensis).
1903. Sowerby. Mar. Invest. S. Afr., u, p. 229 (natalensis).
Aperture longer than spire. Protoconch 2 whorls, diam 1-3 mm., smooth.
Postnatal whorls 5; axial ribs on first 2, sometimes 3 whorls 10, sometimes II
on 3rd, then petering out though occasionally visible on 4th whorl; spiral lirae
4 on each whorl, strong on 4th and 5th whorls, with fine intermediaries on later
whorls, on base 8 additional lirae, with intermediaries; parietal callus denti-
form, columella keeled anteriorly, outer lip plicate within, the uppermost plica
largest and dentiform. 22 X 12 mm.
Orange, with faint white peripheral band. Remains of brown fibrous
periostracum in the spiral grooves.
Living: Umtwalumi and Port Edward (Natal) (U.C.T.).
Dead: off Tugela River (Zululand), 14 fathoms, 1 dead but fresh
(Sowerby) the specimen referred to by Sowerby without locality) (S. Afr. Mus.
PE coll),
Remarks. From the description one suspects that Krauss’s rubiginosus var.
subcostata (1848, Siidafr. Moll., p. 120; quoted in Sowerby. Mar. Sh. S. Afr.,
p- 4, as ‘scabricostata’) was really carinifera.
C. undosus (Linn.), very similar with strong spiral lirae, has also been
recorded (Sowerby 1889, 1892) as var. minor from Port Elizabeth. The locality
is suspect.
Cantharus fumosus (Dillwyn)
1817. Dillwyn. Catal. Sh., p. 629 (Buccinum f.).
1846. Reeve. Conch. Icon., iii, no. 51 (Buccinum proteus).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA I51
1859. Chenu. Man. Conchyl., i, fig. 622 (Buccinum proteus).
1880. Von Martens. Mauritius G Seychellen, p. 239 (Pisania f.).
1911. Schepman. Szboga Exp. monogr., xlix, p. 303.
1957. Robertson. Not. Nat. Philad., 300, figs. 5 (shell), 17 (radula).
A dead but fresh Delagoa Bay specimen was collected by U.W. Postnatal
whorls 5; 10 broad and rounded axial ribs on each whorl, but not extending
across base on body whorl; crossed by spiral lirae 4 on each of the early whorls,
5 on body whorl, rather sharp, with finer intermediaries, 7 additional main
lirae on base, with intermediaries; growth-lines distinct. No parietal callus
(but columella glaze not fully developed), columella keeled anteriorly, outer
lip plicate within. 23 (protoconch missing) x 13 mm. Uniform grey.
Distribution. Mauritius, Seychelles, Madagascar, East Indies.
Remarks. This specimen corresponds well with Chenu’s figure.
Cantharus insculpta (Sow.)
1900. Sowerby. Proc. Mal. Soc., iv, p. 2, pl. 1, fig. 4 (Tritonidea 7.).
Aperture a little longer than spire. Postnatal whorls 5 (Sowerby); axial
ribs 12 on 3rd whorl, 14 on last whorl; crossed by spiral lirae 4 on 3rd whorl,
6-7 on last, 10-11 additional lirae on base; spaces between ribs and lirae rather
deeply pitted on upper whorls, sculpture almost cancellate. Parietal callus
dentiform, columella keeled anteriorly, outer lip smooth within. 11-5 * 6 mm.
(Sowerby).
Yellowish-brown with white peripheral band (at or slightly above the
actual periphery).
Dead: Port Alfred and Kowie (Sowerby, Bartsch, Turton, and S. Afr.
Mus.).
Remarks. 1 have seen only beach-worn specimens lacking the protoconch.
Gen. NassAriA Link
1807. Link. Beschr. Nat. Samml. Univ. Rostock (3), p. 123.
1853. H. & A. Adams. Gen. Rec. Moll., i, p. 123 (Hindsia).
1916. Iredale. Proc. Mal. Soc., xii, p. 82 (Hindsia H. & A. Adams, 1850 [sic]).
1929. Thiele. Handbuch, i, p. 310.
Remarks. Uredale’s argument for adopting Hindsia is not very conclusive;
and Vassaria is not to be rejected on account of the earlier Nassarius (Rule Zool.
Nomencl. Art. 36 Rec.).
Nassaria gracilis Sow.
Fig. 31(4), (¢), (¢)
1902. Sowerby. Mar. Invest. S. Afr., ii, p. 94, pl. 2, fig. 10.
1929. Dautzenberg. Faune Col. Franc., iil, 4, p. 407.
Protoconch 14 (2) whorls, diam. and alt. 0-75 mm., smooth and polished.
Postnatal whorls 84; first half whorl with 6-8 axial ribs, g—1o on the following
152 ANNALS OF THE SOUTH AFRICAN MUSEUM
whorl, increasing to (13) 14 on last whorl, fine growth-lines in the intervening
grooves; crossed by spiral lirae 2 on first two whorls, a third lira appearing
definitely on 3rd whorl but always less prominent than the other two, inter-
mediaries developed from 4th whorl onwards, c. 10 additional lirae on base,
with intermediaries; small complanate nodules at the intersections with the
ribs. Parietal callus dentiform, columella angularly bent, canal narrow and
curved, outer lip plicate within. Periostracum very thin. 29 (protoconch
missing) X 15 mm.
Operculum broadly oval, rather thick and lamellate around margin,
nucleus apical, 5 X 3°5 mm. in 31 mm. shell.
White with pale brown periostracum.
Radula with 75-85 rows, central plate with slightly concave front margin,
8 cusps (6 in another specimen), with 1-2 smaller ones in the middle, the
number of small median cusps varies in different parts of the same radula;
lateral plate bicuspid, outer cusp the larger. A Delagoa Bay specimen (fig.
31(d)) with 85 rows has a total of 9 cusps on the central plate, 5 shorter and
narrower ones flanked on either side by 2 larger and broader ones.
Living: off Tugela River (Zululand), 40 fathoms (Sowerby); off Tugela
River and Amatikulu River, 12-26 fathoms (S. Afr. Mus. P.F. coll.). Delagoa
Bay (U.W.); 30°47’ S. .30° 290 E., 24 fathoms, and 33° 37 S030 sjagee
46 metres (U.C.T.).
Distribution. Madagascar (Dautzenberg).
Remarks. The last rib in shells which appear to be fully adult is larger
than the preceding ones and forms a varix on the outer lip; sometimes a
varicoid rib is developed on one of the previous whorls, the following ribs being
normal; smallest such shell seen: 20 mm. long.
NV. acuminata Rve. has been recorded from Durban (1897. Sowerby. Appen.
Mar. Sh. S. Afr., p. 6), and differs (1902. Sowerby. Mar. Invest. S. Afr., 11, p. 95)
in being larger, proportionately broader, fewer axial ribs, closer and less
conspicuous spiral lirae.
The Delagoa Bay specimens, largest 29 * 16:5 mm., are relatively broader,
but have the same number of axial ribs. The width of course varies according
to whether the shell has reached a stage at which a varicoid outer lip is
developed.
Dautzenberg said the Madagascan example agreed perfectly with a
cotype from mouth of Tugela River, 40 fathoms, in his collection, which came
from the MacAndrew collection. One wonders how a Pieter Faure shell found
its way into the MacAndrew collection!
N. gracilis should be compared also with the Indian nivea (Gmelin) 1790
and suturalis Adams.
Genus AFROCOMINELLA Iredale
1917. Cooke. Proc. Mal. Soc., xii, p. 227 (radula) (Cominella s.1.).
1918. Iredale. ibid., xiii, p. 34, line 22.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 153
1926. Tomlin. Ann. Natal Mus., v, p. 290.
1929. Thiele. Handbuch, i, p. 315 (as a section of Burnupena).
1938. Peile. Proc. Mal. Soc., xxiii, p. 98 (radula).
1944. Stephenson. Ann. Natal Mus., x, p. 344 (Cominella, list of species).
1947. id. ibid., xi, pp. 271-4 (‘Cominella’, distribution and notes on species).
1956. Orr. Proc. Ac. Nat. Sci. Philad., cviii, pp. 249 and 251 (differences from Burnupena,
species not discussed).
Oval-fusiform, spire usually high. Subsutural groove more or less distinct.
Canal moderate. Fasciole absent. Early whorls with clathrate sculpture formed
by spiral lirae and a few (11-12 on first whorl) axial ribs.
Penis without apical prong. Radula central plate as long as broad, with
3 cusps, lateral plate, with strong outer cusp, and bifid inner cusp.
Genotype: elongata (Dnkr.).
Remarks. Thiele united both Iredale’s genera, but he should not have
subordinated Afrocominella, which has line precedence, to Burnupena. |
The retention of two genera depends largely on the value attached to the
differences in the radulae. In adults the two radula are clearly distinct, but in
juvenile Burnupena the central plate may be not so broad as in the adult,
approximating to the squarish shape found in Afrocominella (Peile, 1938, p. 98,
fig. 33; Orr, 1956, p. 261). The lateral plates, however, always serve to
differentiate the two genera. The sculpture of the early whorls seems to provide
an additional distinction.
The two genera are here accepted. Miss Orr has come to the same
conclusion.
Afrocominella contains three species (but not puncturata Sow. which is a
Muricid), which may be distinguished as follows:
1. Axial ribs extending from suture to suture on early and later whorls, but
often feeble on the latter.
a. 7-9 costae on base of last whorl. Only spiral lirae in the subsutural groove
fais? ame Weal Costa) rt wat od <8 dare elise 8a, eA elongata
b. 12-15 costae on base. In the subsutural groove 1-3 well-marked costae capensis
2. Axial ribs on later whorls confined to the periphery, forming shoulder knobs — turtoni
The differences between elongata and capensis are very slight and both are
so variable that it is doubtful whether two species can be maintained; capensis
has priority.
Afrocominella elongata (Dnkr.)
Fig. 31(f)
1857. Dunker. Proc. Zool. Soc. Lond. (for 1856), p. 356 (Cominella e.).
1899. Smith. 7. Conch., ix, p. 248, pl. 5, fig. 3 (Cominella prolongata).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 48, pl. 3, fig. 7 (C. alfredensis).
1917. Cooke. loc. cit., p. 229, fig. 11 (radula) (Cominella e.), and fig. 12 (err.: as tigrina),
and p. 234, fig. 7 (radula, err.: as ‘Euthria queketti).
1918. Iredale. ibid., xiii, p. 34.
1932. Tomlin. Ann. S. Afr. Mus., xxx, p. 166, fig. 7 (Glypteuthria solidissima).
1938. Peile. Proc. Mal. Soc., xxiii, p. 98, fig. 34 (radula).
1938. Eyre and others. Ann. Natal Mus., ix, pp. 96, 109 (Cominella e.).
154 ANNALS OF THE SOUTH AFRICAN MUSEUM
1939. Eyre. ibid., ix, p. 304 (Cominella e.).
1947. Stephenson. ibid., xi, pp. 271-3 (distribution) (Cominella e.).
1952. Day and others. Tr. Roy. Soc. S. Afr., xxxili, p. 410.
1956. Orr. loc. cit., text-fig. 1 2 (radula).
Aperture subequal to spire in juveniles, a little less in larger specimens.
Protoconch 24 whorls, diam. and alt. 1 mm., smooth. Postnatal whorls 9,
whorls with distinct shoulder, spire more or less turreted, especially in early
whorls; 11-12 axial ribs on 1st whorl, increasing to 15-16 (18) on 6th and 7th
whorls, thereafter usually becoming irregular and indistinct, sometimes
becoming obsolete from 5th onwards; crossed by 4 spiral lirae on 1st and 2nd
whorls, the lowest (most anterior) being the strongest, on 3rd and following
whorls the 2 or 3 peripheral lirae are the strongest, being almost costae; from
3rd whorl (at which stage the subsutural groove develops) onwards spiral striae
between the lirae-costae, and especially in the subsutural groove (about 15
between suture and shoulder in later whorls); intersections subnodulose; 7—9
additional costae on base, upper ones more or less subnodulose, with inter-
vening striae. Growth-lines (on all whorls) produce a minutely decussate or
beaded appearance. Parietal callus dentiform; outer lip indented posteriorly
(but not strongly), internally plicate.
61 (protoconch and whorls 1-3 missing) X 24 mm.; 55 (apex missing) X
25 mm.
Operculum oval, scarcely unguiform, 15 < 7 mm. in 52 mm. shell with
aperture 27 mm.
Whitish with irregular chestnut or reddish-brown patches, spots and
flames; periostracum brown; operculum brown. One specimen (Gr. Fish River,
living) without any markings, periostracum grey-brown. Animal speckled
with black or dark grey.
Radula with go-115 rows, central plate about as long (incl. cusps) as
wide (base 14 as wide as long), with 3 cusps, lateral with bifid inner cusp,
inner margin of inner prong feebly and obscurely serrulate. Variations: 4
cusps on central, a triple inner cusp on lateral on one side for part of radula
(see Peile).
Living: South coast from False Bay to Qolora north of East London
(Stephenson 1947); in addition: off Great Fish River, 22 fathoms (S. Afr. Mus.
P.F, ‘coll,). 93°97. 26° 56 EE.) 26 metres) (Gea):
Stephenson (1947) recorded it also from Oudekraal on west coast of Cape
Peninsula, but I have seen these specimens and consider them to be capensis.
Off Cape Point (mouth of False Bay) 45 fathoms (Tomlin: G. solidissima).
Remarks. Sometimes there are three subequal peripheral costae on the
later whorls.
C’. prolongata seems to be merely the full-grown elongata as shown by the
series in S. Afr. Museum. |
Two specimens each of elongata and alfredensis from Port Alfred, collected
and presented by Turton, show no differential characters.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 155
The identity of G. solidissima with A. elongata becomes quite clear when
the details of the sculpture are compared. It might also be compared with the
thick-walled varieties of capensis (p. 156), but only one, perhaps two, of the
spirals in the subsutural groove are strong enough to be called costae.
In this case specimens of elongata in various stages of abrasion were found
useful because the solidissima shell is slightly corroded in places.
Trophon acutispira (Sow.) is very like juvenile elongata of about the same
size, but has fewer costae on the base.
Afrocominella capensis (Dnkr.)
Fig. 31(e)
1844. Dunker in Phillipi. Abd., 1, p. 110, pl. 1, fig. 7 (Fusus capensis).
1852. Petit. 7. Conchyl., iii, p. 164, pl. 7, fig. 7 (Fusus simonianus).
1860. Gould. Proc. Boston Soc., Nat. Hist., vii, p. 327 (Euthrya lacertina).
1874. Von Martens. 7. B. Deutsch. Malak. Ges., i, p. 133, pl. 6, fig. 2 (Euthria lacertina)
(quotes Gould’s description verbatim).
1877. Velain. Arch. Zool. Exp. Gen., vi, p. 104, pl. 2, figs. 8-11 (Euthria magellant).
1903. Smith. Proc. Mal. Soc., V, p. 371 (Euthria c. and lacertina).
1903. Von Martens. D. Tiefsee Exp., vil, p. 25 (Euthria c.).
1925. Thiele. D. Tiefsee Exp., xvii, p. 180, pl. 32 (20), fig. 3 (Pisania costata).
1938. Bright. Tr. Roy. Soc. S. Afr., xxvi, p. 58 (Pollia lacertina).
1939. Peile. Proc. Mal. Soc., xxiii, p. 271 (radula) (E. lacertina).
1947. Stephenson. Ann. Natal Mus., xi, pp. 271-3 (Cominella lacertina).
“~~
Aperture a little larger than spire. Protoconch 24 whorls, diam. and alt.
I mm., smooth. Postnatal whorls 6—7; whorls gently convex, profile of spire
nearly straight, subsutural groove feeble; 11 (12) ribs on 1st whorl, increasing
to 13-14 on 5th, thereafter becoming irregular and indistinct; crossed by 4
spiral lirae on ist and 2nd whorls, 5 on 3rd, lirae becoming costae on later
whorls and increasing in number to 10-12 on last whorl, with 12-15 additional
ones on base; on last whorl costae all approximately of equal strength, but
often a lira between each pair of costae; 1-3 costae in subsutural groove; the
spiral sculpture is usually very regular and clear-cut, with rather deep sulci
between the costae, the sulci striate with closely packed growth-lines. Parietal
callus feeble; outer lip feebly indented posteriorly, internally plicate. 32
(protoconch missing) X 15 mm. }
Operculum almost regularly oval, apex scarcely, if at all incurved,
a4 mm. in 28 mm. shell.
White or greyish with fulvous or chestnut brown spots and axial streaks
and flames, aperture more or less suffused; periostracum greyish-brown;
operculum amber brown. Animal dark grey.
Radula with c. go (juv.) to 110 rows, central plate about as long (including
cusps) as broad, base 14 as wide as long, tricuspid, sometimes with a minute
extra denticle on one side, lateral with bifid inner cusp, inner margin of inner
prong obscurely serrulate.
156 ANNALS OF THE SOUTH AFRICAN MUSEUM
Living: Port Nolloth to Gape Agulhas (Stephenson) ; east and west coasts
of Cape Peninsula (S. Afr. Mus.).
Cape Agulhas (Petit: stmonianus).*
The records (dead shells) from Port Elizabeth (Sowerby), Port Alfred
(Turton), and Natal (Sowerby) need confirmation.
Remarks. A slender specimen, 44 x 18 mm. (S. Afr. Mus. no. 4709)
locality unknown, but probably Cape, was identified by J. H. Ponsonby as
magellan, which Smith (1903) thought might be also synonymous with capensis.
Three dead specimens from Green Point (Cape Town) (S. Afr. Mus.
no. 5518), 20 X 11, 24 X 11 and 25 X 14 mm., white with traces of brown
flames: the smallest is slightly thicker-walled and weighs the same as the 24. mm.
specimen; the largest is plumper and thicker-walled.
Four other dead specimens (S. Afr. Mus. no. 4753), locality unknown,
but probably also Cape Town, 17 X 9, 24 X 12, 26 x 14, and 29 X 15 mm.
are rather plump. The 2 larger ones are thick-walled, the largest especially so,
with a thickened and strongly plicate outer lip. These were identified as
magellan by J. H. Ponsonby.
There are two dead shells, 19 x 10 and 27 x 15 mm., from Table Bay (S. Afr.
Mus. 5465); the larger is thick-walled, and both have the outer lip plicate.
Very different, at first glance, from these thick-walled specimens, are 4
specimens from the P.F. collection but without exact locality (S. Afr. Mus.
A4732). One was taken alive, and the operculum, but not the animal, was
preserved. They are all thin-walled and slender: 25 x 10 and 26 X II mm.
Nevertheless they conform in all characters with the above description.
The ‘straight’ sides, regularity of the spiral sculpture, and the large
number of costae on the base distinguish this species from elongata.
Smith’s suggestion that magellani from St. Paul Island might be synonymous
seems rather unlikely, but should be investigated. Sowerby’s record (1897) of
magellam from Natal is certainly erroneous.
In spite of Thiele’s statement that the Valdivia specimen is different in
shape from Euthria capensis, it might perhaps be assigned to this species or to
elongata. Thiele described it (or another specimen) as Prsania costata. ‘There is
a specimen from Sea Point (Cape Town) (S. Afr. Mus. no. 4974) of almost the
same size (Thiele 18 7:5 mm., S. Afr. Mus. 18 x 8 mm.); if Thiele’s figure
correctly indicates the size of the costae in the subsutural groove, and the
number of costae on the base, Pisania costata can well be regarded as a
capensis. ‘The axial ribs, however, appear from the figure (the number is not
given in the description) to be fewer. Von Martens’s and Thiele’s locality:
94° 51'S, 19° 37 E., 80 metres.
There is little difference in Petit’s figure of his simonianus and von Martens’s
figure of lacertina. I think there is little doubt that szmonianus should fall into
the synonymy of capensis.
* Named after M. de Saint-Simon, not after Simon’s Bay.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 157
The difficulty of assigning some specimens to a particular species is shown
by the following description of a specimen labelled by Sowerby (3rd) as
‘simoniana juv.’ with a query.
Fusiform, profile of whorls very slightly convex. Protoconch 24 whorls,
diam. and alt. 1 mm., smooth. Postnatal whorls 5; low axial ribs 13 on Ist
whorl, increasing to 18 on last, on 1st—3rd whorls distinct from suture to suture,
but becoming indistinct on 4th and first part of 5th and indicated only by a
double row of feeble peripheral knobs, and obsolete on body whorl; crossed by
very fine spiral striae, visible chiefly between suture and periphery on last two
whorls and on base, on the latter there are 4 or 5 slightly stronger lirae, and
half a dozen closer together on rostrum. No parietal callus, columella curved,
canal rather narrow and reflexed, outer lip srmple. 21 X 9 mm. Pale buff.
Off East London, 40 fathoms (S. Afr. Mus. P.F. coll.).
Remarks. This specimen is closely similar to a 22 X Io mm. specimen of
Afrocominella elongata from Qolora, and also with some of the specimens (A4732)
referred to A. capensis, though much smoother.
Afrocominella turtont (Bartsch)
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 50, pl. 3, fig. 6 (Euthria t.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 54 (Euthria t.).
Spire turreted. Profile with shoulder at middle of whorl. Protoconch 24
whorls, diam. 1-2, alt. 1 mm., smooth. Postnatal whorls 5; axial ribs 10 on
Ist and 10-11 on 2nd whorl, increasing to 12 on 5th whorl, from suture to
suture on Ist and 2nd, but from 3rd whorl onwards evanescent between suture
and periphery, where they form prominent shoulder knobs; not extending
across base; crossed by 4—5 fine spiral lirae on 1st whorl, on 2nd whorl 4 above
and 3 below the (incipient) shoulder, increasing to 12-15 above and 10-12
below on 5th whorl; base with c. 30 additional closely packed lirae, of which
7 or 8 are slightly stronger than the others. Parietal callus not developed, canal
straight, outer lip not plicate within. 22 X 9:5 mm.
Operculum oval, nucleus apical, scarcely incurved, 5 X 2°5 mm.-in
22 mm. shell with 11 mm. aperture.
Castaneous above the shoulder, and between the knobs, ochraceous below,
operculum amber. Animal pale with dark grey speckling around foot.
Radula with c. 120 rows, central plate about as long (incl. cusps) as wide,
base rectangular, with 3 cusps, lateral plate with bifid inner cusp.
St. Francis Bay (34° 15’ 8. 25° 5’ E.), 6 fathoms (U.C.T.).
Remarks. The above description is taken from the, as yet, only specimen
known to have been collected alive. Its radula showed it to be an Afrocominella.
This was not unexpected because Turton said (loc. cit., p. 52) that some of his
specimens were identified by Smith as capensis, and S. Afr. Mus. specimens
with the turton coloration were identified many years ago by J. H. Ponsonby as
elongata.
158 ANNALS OF THE SOUTH AFRICAN MUSEUM
Beach-worn examples from Port Alfred and Still Bay (S. Afr. Mus.) up to
40 mm. in length, with 6-7 whorls, yellowish or orange-brown and showing
more or less the castaneous patches between the knobs, agree in sculpture with
the above description. A second row of less prominent nodules below the
shoulder knobs is sometimes present. The parietal callus is nodiform.
Three P.F. specimens, dirty white without any colouring, also agree, but
are thicker walled, especially the largest. The outer lip tends to be slightly
exsert and thickened within, with about 1o denticles in the largest specimen
(merely indicated in the other two).
33 (apex and rostrum broken) xX 17 mm. (7 whorls); 26 (apex and tip of
rostrum broken) X 12 mm. (6 whorls); 17°5 <x 8 mm. (5 whorls).
Off Great Fish Point, 57 fathoms (largest); off East London, 32 fathoms;
off Hangberg (Knysna, not Hangklip, False Bay), 48 fathoms (smallest); one
dead but fresh example from each locality (S. Afr. Mus. P.F. coll.).
Gen. BURNUPENA Iredale
1917. Cooke. Proc. Mal. Soc., xii, p. 227 (radulae) (Cominella s.1.).
1918. Iredale. ibid., xii, p. 34, line 24.
1926. Tomlin. Ann. Natal Mus., v, p. 291.
1929. Thiele. Handbuch, i, p. 315 (incl. Afrocominella).
1938. Bokenham & Neugebauer. Ann. Natal Mus., ix, p. 133 (egg-capsule).
1938. Peile. Proc. Mal. Soc., xxiii, p. 97, and 1939, p. 270 (radulae).
1944. Stephenson. Ann. Natal Mus., x, p. 344 (list of species, Cominella).
1947. id. ibid., xi, pp. 271-4 (distribution and notes on species, Cominella).
1951. Barnard. Beginner’s Guide S. Afr. Shells, fig. 35 (egg-capsule).
1956. Orr. Proc. Ac. Nat. Sci. Philad., cviii, p. 250 et sqq. (revision and ecology of species) .*
Ovate, sometimes slightly fusiform, spire low or not very high. Subsutural
canal usually distinct. Canal short. Fasciole distinct. Early whorls either
cancellate-nodulose (axial ribs on ist whorl numerous: 18-20), or with spiral
sculpture only.
Penis with apical prong. Radula central plate wider than long, with
(3-4 juv.) 5-7 (8) cusps, lateral plate with strong (usually not so strong as in
Afrocominella) outer cusp and trifid inner cusp. Malformations and asymmetry
frequent.
Genotype: cincta (Bolten-Réding).
Remarks. The egg-capsule and protoconch described below undoubtedly
belong to a Burnupena, but whether the species is cencta or papyracea is uncertain.
I have found them at St. James (False Bay), and Bokenham and Neugebauer
collected them at Sea Point (Cape Town) and St. James. Actual spawning
has not been observed.
After examining the S. Afr. Mus. collection (mostly dead specimens), and
the large U.C.T. collection of material taken alive by Prof. Stephenson and
Prof. Day, I have come to the conclusion that six species: cincta, lagenaria,
* Some typ. err. and laps. cal. In the Bibliography, Bokenham & Neugebauer 1938, for
‘Tr. Roy. Soc. S. Afr.’ read: Ann. Natal Mus.; for ‘Piele’ read: Peile.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 159
papyracea, limbosa, delalandit and tigrina, can be more or less satisfactorily
separated.
Stephenson’s remarks (1947, p. 272), however, are fully justified. Some
examples are impossible to assign definitely to one species or another, owing
to the frequent occurrence of intergrading forms which Stephenson suggested
might possibly be hybrids. The two most confusing pairs are cincta-lagenaria
and limbosa-delalandu. But in spite of intergrading forms, the extremes are
distinctive, and for this reason might well be retained as separate species.
A. Early whorls with spiral sculpture only.
1. Whorls with spiral lirae and more or less numerous well-marked costae.
Pale spots around spire (if not corroded).
Pee WEP CTOMNS INC NGO Omni eR nt cette fo kee mn ae ay! 6 | ely ge |e 5 CONCID
(intermediates)
MAH UCIOUS ITC FON Oh) Foe 2 | fe te hes, Sele «he on, 9 Cagenarta
2. Numerous striae and lirae, but none enlarged to costae.
a. Subsutural groove not, or only feebly, developed. Profile of whorls
Suen COnvexs MPerEkenvinite Vey 2t i os a ls AE Sey Se papyracea
(intermediates)
b. Subsutural groove well developed.
muBrown, aperture more) or lesssuffused . <7). <3. |... \.. lambosa
(intermediates)
ii. Purplish-brown, aperture deeply suffused purplish-brown or
OAC OMS NES Steere ee Nhe aM Maa uae a= 2s cin pedelaanare
B. Early whorls cancellate-nodulose, the axial ribs numerous (18-20) . . . tigrina
To Stephenson’s summaries of the distribution of each species are added
the localities of the S. Afr. Mus. examples (when definitely known). Little
reliance, however, can be placed on some of the earlier records.
Only selected references to each species are given, mainly those of recent
years. In some cases likely synonyms are suggested.
Since the completion of my study of this genus, Miss Orr’s valuable paper
has appeared. It is refreshing to have a revision of a genus based on living
material personally collected and studied in the field. After investigating
habitats and ecology at several localities around the South African coast,
Miss Orr has reduced the number of species to two; papyracea and delalandii,
the former with papyracea s.s., cincta, lagenaria and tigrina as subspecies.
To some extent shell characters were found correlated with habitat, e.g.
the strongly costate cincta inhabits intertidal pools along the south coast,
finely striate form s:being found in the colder subtidal zone, and especially on
the west coast; shells in sheltered situations usually have higher spires than
those from exposed habitats.
Basically there is little difference between Miss Orr’s conclusions and my
own; the main difference concerns the taxonomic status of the forms. Feeling
that the last word, ecologically and taxonomically, has not been said, I am
retaining my diagnoses.
The spawning habits need investigation. And if in the future it should be
found possible to breed these molluscs under artificially controlled conditions,
~ a
oy :
160 ANNALS OF THE SOUTH AFRICAN MUSEUM
instructive results may be obtained. It would be interesting, for example, to
see what sculpture would be found on the progeny of a cincta x tigrina hybrid
(cf. fig. 32(e) and (/)).
Burnupena cincta (Bolten)
Figs. 31(g), 32 (a-d), (e)
1790. Gmelin. Syst. Nat., ed. 13, p. 3494, no. 105 (Buccinum porcatum, non da Costa 1778).
1798. Bolten-Réding. Mus. Bolten., p. 113.
1874. Von Martens. J.B. D. Malak. Ges., p. 136 (Buccinum p.).
1886. Watson. Challenger Rep., xv, p. 214 (Cominella p.) (references).
1889. Studer. Forschungsreise d. ‘Gazelle’, 111, pp. 52, 55 (Buccinum p.).
1903. Von Martens. D. Tiefsee Exp., vii, p. 52 (Cominella p.).
1917. Cooke. loc. cit., p. 229, fig. 1 b (radula) (Cominella p.).
1918. Iredale. ibid., xiii, p. 34.
1926. Tomlin. Ann. Natal. Mus., v. p. 291.
1932. Turton. Mar. Sh. Pt. Alfred, p. 52, pl. 12, no. 381 (cincta var. adjacens).
1937. Stephenson and others. Tr. Roy. Soc. S. Afr., xxvi, p. 357.
1938. Peile. Proc. Mal. Soc., xxiii, p. 97 and p. 98 (dunkeri (radula).
1938. Stephenson. Ann. Natal Mus., ix, p. 10.
1938. Eyre and others. ibid., ix, p. 96.
1938. Bokenham & Neugebauer. ibid., ix, p. 133, pl. 16, figs. 1-3, 5, 6, 9 (egg-capsule,
development, protoconch).
1939. Eyre. ibid., ix, p. 298.
1940. Broekhuysen. Tr. Roy. Soc. S. Afr., xxvili, pp. 255 et passim (fig. 1 chart of vertical
distribution).
1947. Stephenson. Ann. Natal Mus., xi, pp. 272, 273.
1952. Day and others. Tr. Roy. Soc. S. Afr., xxxiil, p. 410.
1956. Orr. loc. cit., p. 254, pl. 19, figs. 5, 6, text-fig. 1 b, c, pl. 20, fig. 2.
No axial sculpture on any of the whorls. Aperture a little longer than
spire. Ratio breadth to length = 1 : 1-6-2. Angle of spire 50°-65°. Proto-
conch ovoid, 24 whorls, diam. 1-3—1-5 mm., alt. 2-2-5 mm.* (specimens taken
from egg capsules), smooth but with c. 15 spiral lirae beginning on last half
whorl. Postnatal whorls 7, with subsutural groove; 3 broad flattish spiral costae,
the uppermost adjoining suture, the 2nd forming lower border of subsutural
groove, the 3rd usually partly or wholly covered on the upper whorls by the
succeeding whorl; 3 (nos. 4, 5, 6 in fig. 32(a)—(d)) additional costae on base of
body whorl; usually an additional narrower costa between each pair of major
costae, but not between Ist and 2nd (except in one specimen); costae and
grooves (incl. subsutural groove) with fine spiral striae. Parietal callus denti-
form; canal reflexed; outer lip undulate on margin (corresponding with the
costae), but usually not plicate internally (in fresh intact specimens the actual
edge may be finely crenulate, corresponding with the striae). Periostracum
thick, fibrous-fimbriate. 59 (apex corroded) x 31 mm. Plump and slender
forms, e.g. 50 X 30 mm. and 51 X 27 mm. (same locality).
Operculum ovate, somewhat unguiform, apex incurved, 23 X II mm.
in 55 mm. shell.
* i.e. including the sculptured portion—smooth portion o-g-1 mm.
10
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 161
Orange-brown with darker marks and spots alternating with white marks
on the major costae; coloration obscured in living examples by dark brown
periostracum, except on the upper whorls where the alternate brown and white
‘necklace’ remains on Ist and 2nd costae; in large specimens the early whorls
are nearly always corroded; aperture and columella white, but outer lip some-
times slightly suffused internally; operculum dark brown. First whorl of
protoconch white, second fawn. Periostracum often stained with green algae.
Animal mottled or streaked with blackish-grey.
Radula with 110-120 rows, central plate wider than long, with 6 denticles
(sometimes only 5), lateral with trifid inner cusp, the 2 inner prongs feebly
serrulate on their opposing edges. Peile: denticles on central 5-9, usually 6-7,
his fig. 33 of dunkeri is a juvenile radula with central plate not so wide as in
adult and with only 3 cusps.
Egg-capsule a triquetral prism, length 7-9 mm., maj. diam. 3—3°5 mm.;
outer (convex) surface with a median keel in the basal quarter or third; apex
sometimes fimbriate. Numerous capsules are attached by their base in oval
or circular clumps, }” to 2” in diameter, to rock-surfaces or fronds of seaweeds.
Living: False Bay to The Haven (north of East London), and sporadically
on west coast as far north as Steenberg Cove (St. Helena Bay) (Stephenson
1947); also Richmond (Alexandria Division) (U.C.T. Ecol. Surv.); the west
coast localities are Sea Point (Cape Town), Langebaan, Steenberg Cove, and
Liideritzbucht (U.C.T.). Simon’s Bay (False Bay) 10-20 fathoms (Watson).
Kalk Bay and Zwartklip in False Bay (S. Afr. Mus. coll. K.H.B.).
Dead: several localities within the above range (previous authors, and
S. Afr. Mus.). 34° 6’S. 18° 6’ E., 117 fathoms (Studer), Saldanha Bay, and
other localities to Natal (von Martens), Cape Congo, Angola (Orr).
Remarks. As a rule the spiral costae are well developed and broad; where
intermediaries are developed the major costae tend to be narrower. On the
body whorl there is a possible maximum of 10: 6 major and 4 intermediaries
(fig. 32(a)); a total of 7, 8 or 9 is common, but 10 is rare. Sometimes nos. 2, 3
and intermediary 2a are subequally narrow (fig. 32(b)); in another variation
all the costae may be narrow and consequently widely spaced (fig. 32(c)).
In only one specimen have I seen an intermediary in the subsutural
groove between the ist and 2nd major costae, thus tending to obliterate the
groove (fig. 32(d)).
The costae are sometimes very flat.
There are high-spired (czncta s.s.) and low-spired forms, the latter being
broader, more squat, and approaching the lagenaria shape (cf. Stephenson
1947). The U.C.T. collection has transitional examples from Langebaan (West
Coast), St. James (False Bay), Danger Point, Cape Agulhas, Mossel Bay,
Kleinmond and Richmond (Bathurst and Alexandria Div.), Qolora, Port St.
Johns.
From Liideritzbucht (U.C.T. L.U. 29.C.) I have seen two specimens of
162 ANNALS OF THE SOUTH AFRICAN MUSEUM
somewhat squat form, brightly coloured with flames in the subsutural groove,
and ‘necklaces’ of dark and light spots on each costa.
One specimen from Kalk Bay (False Bay) with aberrant operculum:
oval with subcentral nucleus (cf. lagenaria aberr.).
Burnupena lagenaria (Lam.)
Fig. 31(h)
1822. Lamarck. Anim. sans Vert., vil, p. 245 (Purpura I.).
1832. Duclos. Ann. Sci. Nat., xxvi, p. 112, pl. 2, fig. 12 (P. cucurbita).
1903. Von Martens. D. Tiefsee Exp., vii, p. 52 (Cominella l.).
1910. Schwarz. Tr. Geol. Soc. S. Afr.; xii, p. 114.
1917. Cooke. Proc. Mal. Soc., xii, p. 229, fig. 14 (radula).
1922. Tomlin. 7. Conch., xvi, p. 260.
1929. Thiele. Handbuch, i, fig. 349 (radula).
1938. Peile. Proc. Mal. Soc., xxiii, p. 97 (radula).
1938. Stephenson and others. Ann. Natal Mus., ix, p. 10.
1938. Eyre and others. ibid., p. 96.
1938. Eyre. ibid., p. 298.
1947. Stephenson, ibid., x1, pp. 272-4.
1956. Orr. loc. cit., p. 256, pl. 19, fig. 9, text-fig. 1 g, h.
No axial sculpture on any of the whorls. Proportionately broader than
cincta: ratio breadth/length 1: 1-3~-1-6; angle of spire 70°-85°. Fine spiral lirae
on all whorls. Spiral costae as in cincta, i.e. 6 major costae with a varying
number of intermediaries (1-4), but much flatter and less well developed;
nevertheless, the major costae, though faint, can be distinguished from the
finer lirae. Parietal callus dentiform; outer lip more often plicate on margin
but the plicae not extending internally. Upper whorls usually corroded.
40 X 26 mm.
Operculum ovate, apex somewhat incurved, 12 xX 8 mm. in 30 mm.
shell, 14 X 8 in 32 mm. shell.
Coloration similar to that of cincta: in worn shells the ‘necklace’ pattern is
well marked, but not so obvious when covered by the periostracum; outer lip
internally strongly suffused with orange-brown or chestnut, often with a livery
tinge (Krauss: “‘brownish-violet’), paler at the actual margin.
Radula with 120-130 rows, central plate wider than long, with 6(7)
denticles (Thiele’s figure shows 5 denticles with a minute 6th at one side;
Cooke shows 6 plus 1; Peile gives 4-8, 5 being the normal number); lateral
with trifid inner cusp.
Fossil: Pleistocene, Port Elizabeth (Schwarz).
Living: False Bay to Umbhlali, Natal (Stephenson 1947); Zwartklip,
False Bay (S. Afr. Mus. coll. K.H.B.).
Dead: various localities within the above range (previous authors) ; also
Walfish Bay (von Martens); Liideritzbucht (von Martens, Tomlin); Table
Bay (von Martens).
Remarks. The broad squat form with deeply suffused aperture is easily
distinguished from the high-spired cincta with pale aperture. But there are
intermediates combining a high spire with a suffused aperture.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 163
Moreover, juveniles are often particularly difficult to assign to one or the
other species. See Stephenson 1947, and examples in U.C.T. Ecol. Surv. Coll.
Aberrations. One freak subscalariform specimen taken alive in False Bay
(S. Afr. Mus. 11117), 27 mm. long, has the spire 13 mm., width 14 mm.
aperture 14 mm. long, (ratio breadth/length almost 1:2); angle of spire
approx. 40°. Almost the whole shell is badly corroded, but the uncorroded
part (¢. 1o mm.) of the last whorl retains the periostracum and shows 6 very
flat major costae, with intermediaries anteriorly; aperture internally suffused.
Identified by Tomlin as lagenaria; I agree.
In one specimen the operculum is oval with nucleus subcentral. (cf. cincta
aberr.).
Burnupena papyracea (Brug.)
1789. Bruguiére. Encycl. Meth. Vers., 1, p. 260 (Buccinum p.).
1816. Lamarck. Tabl. Encycl., pl. 400, figs. 3 a, 3 b, and Liste, p. 2 (Buccinum p.).
1846. Reeve. Conch. Icon., pl. 5, sp. 32 (Buccinum intinctum).
1848. Krauss. Siidafr. Moll., p. 120 (Buccinum intinctum).
1903. Von Martens. D. Tiefsee Exp., vii, p. 52 (Cominella p.).
1922. Tomlin. 7. Conch., xvi, p. 260.
1938. Bright. Tvans. Roy. Soc. S. Afr., xxvi, pp. 62, 72.
1939. Peile. Proc. Mal. Soc., xxiii, p. 270.
1940. Stephenson and others. Ann. Natal Mus., ix, p. 356.
1947. Stephenson. ibid., xi, pp. 273, 274.
1956. Orr. loc. cit., p. 252, pl. 19, figs. 1-4, text-fig. 1 d, e.
Profile of whorls convex, without subsutural groove except sometimes a
trace on last whorl in adult. Protoconch 24 whorls, diam, and alt. 1-5 mm.,
smooth. Postnatal whorls 5; spiral lirae 5 on 1st whorl, increasing to 14-15
on last whorl, with c. 20 additional on base; on early whorls the lirae regular
in size and spacing, but on base the alternating intermediaries are weaker.
Parietal callus dentiform; outer lip very feebly indented posteriorly in adult,
internally plicate. 45 (apex broken) x 26 mm.; 57 (apex and lower part of
outer lip worn) X 32 mm.
Operculum ovate, apex somewhat incurved, 15 X 7 mm. in 40 mm. shell.
Uniform yellow, orange-brown or castaneous, aperture internally white
(except in thin-walled juveniles), periostracum and operculum brown.
Radula with 115 (one specimen examined; Peile 95-100) rows, central
plate with 6 denticles, lateral with trifid inner cusp, the 2 inner points with
their opposing margins feebly serrulate.
Living: west coast from Port Nolloth to Cape Peninsula, and eastwards
as far as Hermanus (Stephenson 1947); Liideritzbucht (U.C.T.).
Dead: Still Bay and Port St. Johns (S. Afr. Mus.); Liideritzbucht (von
Martens, Tomlin); Olifants River, Table Bay, False Bay, Pondoland (von
Martens); other records by previous authors, extending to Natal, require
confirmation.
Distribution. Gabun, French Equatorial Africa (Orr).
164 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. Description based on specimens (S. Afr. Mus.) identified by
Tomlin, who used the name papyracea. Krauss preferred to regard the South
African specimens as a species different from the Norwegian (fide Kiener)
papyracea under the name intincta Rve.
The uppermost 4 lirae on the penultimate and ultimate whorls occupy
the space of the undeveloped subsutural groove; the 4th may be a trifle stronger
and form a very obscure shoulder (but without a concave groove above). The
convex, in plump examples almost globose, profile of the whorls is the most
noticeable feature. The white aperture also seems characteristic.
The plicae within the aperture are already developed in juveniles 6—7 mm.
long.
Two specimens in the Preter Faure collection (S. Afr. Mus. no. A8607.
P.F. 12558) come from: Cape Natal W x N 2? N. 185-200 fathoms. The depth
seems excessive for species of this genus, and probably the specimens have been
mislabelled.
Frequently covered with a dark purplish-brown Polyzoan (Alcyonidium
nodosum), which is not known to occur on any other shell (Stephenson, 1947,
p- 273). I have examined the U.C.T. Ecol. Survey collection and can confirm
Stephenson’s statement. Whenever limbosa and papyracea occurred at the same
locality, specimens with the Polyzoan coat proved to be papyracea as here
diagnosed.
Specimens of tigrina, however, were also found to be covered with a very
similar growth, which has not yet been identified.
Burnupena limbosa (Lam.)
1822. Lamarck. Anim. sans Vert., vii, p. 243 (Purpura l.).
1903. Von Martens. D. Tiefsee Exp., vii, p. 52 (Cominella l.).
1908.* Melvill & Standen. Tr. Roy. Soc. Edinb., xlvi, 1, p. 154.
1909. id. Sci. Res. ‘Scotia’, v, p. 124 (reprint of previous paper).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 47, pl. 4, fig. 6 (C. porcata var. multilirata).
1917. Cooke. Proc. Mal. Soc., xii, p. 229, fig. 15 (radula) (Cominella 1.).
1923. Odhner. Géteb. K. Vet. Handl., xxvi, p. 6 (Cominella 1.).
1926. Tomlin. Ann. Natal Mus., v, p. 291.
1938. Bright. Tr. Roy. Soc. S. Afr., xxvi, pp. 58, 62.
1938. Peile. Proc. Mal. Soc., xxiii, p. 97 (radula).
1939. id. ibid., xxiii, p. 270, fig. 38 (radula).
1940. Stephenson and others. Ann. Natal Mus., ix, p. 356.
1947. Stephenson. ibid., xi, pp. 273, 274.
1956. Orr. loc. cit., p. 252 (as syn. of papyracea).
Ratio breadth/length 1: 1-6-1-8. Angle of spire 70°-g0°. Aperture twice
spire. Protoconch ? Postnatal whorls ? 5. Subsutural groove present, but no
sutural costa. Whole whorl with numerous spiral lirae of almost equal strength,
but those on base somewhat stronger. Parietal callus dentiform; canal reflexed;
outer lip with numerous plicae extending inwards from the margin. Perio-
* Issued separately 1907.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 165
stracum thick, fibrous-fimbriate. 60 x38 mm. Also: 58 x 31 mm. (apex
and lower part of outer lip worn), 56 x 33 mm.
Operculum ovate, apex somewhat incurved, 17 X 9 mm. in shell
Av xX 26 mm.
Uniform yellowish-brown, aperture more or less suffused with fawn at
margin internally, periostracum brown, operculum dark brown.
Radula with c. 130 rows, central plate with 6 denticles (Peile gives 6-8,
usually 7), lateral with trifid inner cusp. Peile (1939) figures a variation.
Living: west coast from Port Nolloth to the Cape Peninsula, further east ?
(Stephenson 1947).
Dead: Table Bay (Sowerby, and S. Afr. Mus.); Walfish Bay, Liideritz-
bucht, Saldanha Bay, Table Bay (von Martens); other records by previous
authors require confirmation.
Remarks. Distinguished from lagenaria by the uniform strength of the
lirae, without any suggestion of major costae; and also by coloration.
Why multilirata was made a subspecies of porcata (= cincta) was not
explained by Bartsch; it appears to be a normal specimen of limbosa.
Orr regards limbosa as a synonym, not even a subspecies, of papyracea
papyracea (p. 253).
Burnupena delalandu (Wiener)
1833. Quoy & Gaimard. Voy. Astrolabe Moll., p. 456, pl. 30, figs. 32-34 (Buccinum violaceum).
1834. Kiener. Cog. Viv., p. 33, pl. 8, fig. 23 (B. violaceum).
1834. id. ibid., ix, p. 15, pl. 5, fig. 14 (Buccinum delalandit).
1848. Krauss. Stidafrik. Moll., p. 120 (Buccinum v. and d.).
1917. Cooke. Proc. Mal. Soc., xii, p. 229, fig. 13 (radula) (Cominella d.).
1918. Iredale. ibid., xiii, p. 34 (Burnupena delalandit).
1923. Odhner. Géteb. K. Vet. Sam. Handl., xxvi, p. 6 (Cominella d.).
1926. Tomlin. Ann. Natal Mus., v, p. 291 (Afrocominella d.).
1938. Peile. Proc. Mal. Soc., xxiii, p. 97, fig. 31 (radula).
1938. Bright. Tr. Roy. Soc. S. Afr., xxvi, pp. 76, 84, 86, 87 (Cominella d.).
1940. Stephenson and others. Ann. Natal. Mus., ix, p. 356 (Cominella d.).
1947. Stephenson. ibid., xi, pp. 272, 273 (Cominella d.).
1956. Orr. loc. cit., p. 258, pl. 19, fig. 10, text-fig. 1 f, pl. 20, fig. 1 (delalandit).
Aperture a little larger than spire. Protoconch 24 whorls. Postnatal
whorls 5; subsutural groove present; spiral striae on all whorls, about 8 on 3rd
whorl increasing to 15-20 on later whorls, with c. 16 additional ones on base,
the latter becoming slightly deeper and farther apart anteriorly (more like
shallow sulci). Parietal callus dentiform; outer lip indented posteriorly,
posterior canal well marked; internally plicate. Periostracum fibrous-fimbriate.
55 (apex corroded) xX 30 mm.
Operculum ovate, somewhat curved inwards apically; 20 x 10 mm. in
55 mm. shell.
General coloration dark purplish-brown, especially juveniles, larger
examples show purplish lirae on a paler ground colour on the base and where
the periostracum is worn; sometimes with dark axial flames on the paler
166 ANNALS OF THE SOUTH AFRICAN MUSEUM
ground, but these more often visible in dead and worn specimens than living;
periostracum brown; aperture violaceous or purplish-brown, operculum dark
brown.
Also, flames appear to be more frequently developed in some localities
e.g. Langebaan Lagoon and Hondeklip Bay.
Radula with c. 110-120 rows; central plate wider than long, 6—7 denticles
(Peile gives 4—7, usually 6), lateral with trifid inner cusp, the inner and middle
points of which are feebly serrulate on their opposing margins.
Living: Stephenson (1947) records this species as common on the west
coast from Port Nolloth southwards, and extending on the south coast as far
as Hermanus.
In view of Stephenson’s results, based on living material, the records of
Bartsch and Turton (6 specimens and 1 resp.) which are the only records east
of Cape Agulhas, can be disregarded.
Dead: (violacea) ‘Table Bay (Quoy & Gainard); (delalandi) Cape (Kiener) ;
Dyer Island (Agulhas) (Odhner). Kalk Bay (False Bay), Dassen Island
(Table Bay), (S. Afr. Mus.).
Remarks. The dark purplish colour with violaceous aperture of living
examples is distinctive, and suggests that this was the species collected in
Table Bay and described by Quoy & Gaimard as violacea. However, as I have
not seen the original figures, or the specimen (if it is still extant!), and as there
are other possible shells (e.g. Thais capensis, T. dubia, perhaps even a worn and
stained Fasciolaria lugubris), Kiener’s name is retained. Orr regards violacea as
a synonym of papyracea cincta.
The apices of all specimens I have seen are more or less corroded; con-
sequently the details of the protoconch and first two whorls cannot be given.
Smallest specimen seen 8-5 mm. long.
There are slender and plump examples, irrespective of sex, e.g.:
Go 20 10, 64% 21 mime:
29 30 X 18, 33 X 21, 34 X 22, up to 53 X 32, 55 X 30 mm.
There are no clear-cut conchological differences between this species and
limbosa. Fresh specimens may be separated by coloration, but worn and faded
individuals are impossible to assign to one or the other ‘species’.
Burnupena tigrina (Kien.)
Fig. 32(/)
1834. Kiener. Cog. Viv., ix, p. 27, pl. 10, fig. 32 (Buccinum t.).
1848. Krauss. Siidafr. Moll., p. 120 (Buccinum t.) (says name is preocc. by Gmelin and
must be changed).
1892. Sowerby. Mar. Sh. S. Afr., p. 10, pl. 1, fig. 7 (Cominella semisulcata).
1917. Cooke. Proc. Mal. Soc., xii, p. 229, fig. 12 (radula) (Cominella t.).
1918. Iredale. ibid., xiii, p. 34.
1932. Turton. Mar. Sh. Pt. Alfred, p. 51, pl. 12, no. 379 (Cominella translucida).
1938. Peile. Proc. Mal. Soc., xxiii, p. 97, fig. 32 (radula).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 167
1947.. Stephenson. Ann. Natal Mus., xi, pp. 273, 274.
1956. Orr. loc. cit., p. 255, pl. 19, figs. 7, 8, text-fig. 1 a.
Not 1917. Cooke. Proc. Mal. Soc., xii, p. 229, fig. 12 (radula) (= A. elongata).
Aperture longer than spire. Protoconch 24 whorls, diam. 1:3 mm.,
smooth but with nodulose-cancellate sculpture in last half whorl. Postnatal
whorls 6; early whorls cancellate-nodulose; 18 (19) riblets on 1st whorl,
increasing to c. 28-30 on last whorl, extending from suture to suture except on
5th and 6th whorls where they form only nodules on the subsutural costa and
the costa forming lower-border of subsutural groove, and less distinctly on
some of the costae below; spiral costae 4 (sometimes 5), of equal strength on
Ist and 2nd whorls, but on 3rd and later whorls the subsutural costa and the
one forming the shoulder become stronger than the others; also from 3rd
whorl onwards an intermediary develops in the subsutural groove, increasing
to 3 or 4 on last whorl; 9-10 additional costae on base; all the costae and
intervening grooves may have fine lirae. Parietal callus dentiform; outer lip
indented posteriorly, internally smooth (lirae showing through half-grown
shells simulate plicae, but this is due to colour only, not sculpture). Perio-
stracum fibrous. 40 X 22 mm., 41 X 20 mm.
Fic. 32.
(a)—(d) Burnupena cincta (Bolten) body whorl to show variation of costae (semi-diagrammatic).
(e) protoconch. (/) B. tigrina (Kien.) protoconch.
168 ANNALS OF THE SOUTH AFRICAN MUSEUM
Operculum ovate, somewhat incurved apically, 8-5 x 4 in 25 mm. shell.
‘Leonine’ coloration: uniform tawny or yellowish-brown, periostracum a
little darker.
‘Tigrine’ coloration: with chestnut or reddish-brown spots and marks on
the costae forming irregular, more or less disconnected axial undulate or
zigzag flames; very obvious in beach-worn specimens, but more or less obscured
by the periostracum in fresh specimens.
Radula with 100-115 rows, central plate twice as broad as long, with 5-6
denticles, the 6th being sometimes minute, lateral with trifid inner cusp, the
opposing margins of the 2 inner points feebly serrulate. (Cooke’s description
and figure incorrect—see Peile 1938.)
Living: Still Bay and East London (Stephenson 1947); False Bay, Her-
manus, Mossel Bay, Jeffreys Bay, Port Elizabeth (U.C.T. Ecol. Surv.); False
Bay, 9 fathoms, and off Cape St. Blaize, 17 fathoms (S. Afr. Mus. P.F. coll.);
Saldanha Bay (west coast) (U.Q.T.).
Dead: records by previous authors are within the above range. Kalk Bay
and Muizenberg (False Bay), Still Bay, Mossel Bay, Port St. Johns, Cove Rock
(near East London), and Durban (S. Afr. Mus.); on the west coast: Dassen
Island (Table Bay), and Lambert’s Bay (S. Afr. Mus.).
Remarks. The above description based on 3 specimens with the ‘leonine’
coloration from False Bay, one of them identified by Tomlin.
There are plump and slender forms: e.g. 40 X 22mm., and 41 X 20mm.;
the uniformly coloured ‘leonine’ examples are mostly plump, the ‘tigrine’
examples on the other hand tend to be more slender.
Some specimens have faint indications of spots and flames and constitute
transitions between the ‘leonine’ and ‘tigrine’ forms.
Sowerby had one specimen, 50 X 24 mm., for his description of semi-
sulcata. His figure is not good, because the subsutural groove is drawn too deep,
simulating a sunken suture, whereas the actual suture lies above the upper
series of nodules. There appear to be no later records of this form.
Five beach-worn specimens (S. Afr. Mus. Ross-Frames coll.), however,
seem referable. On the 4th and 5th whorls the gradually disappearing axial
ribs give a beaded appearance to the subsutural costa, and less obviously to
the shoulder below the groove; but eventually they disappear completely. The
sculpture of the early whorls resembles that of tigrina. 56 (apex and end of
rostrum worn) X 24 mm.; smallest 25 x 13 mm.
Variation. In four ‘leonine’ specimens from Muizenberg (False Bay)
(S. Afr. Mus. no. A4936), largest 38 mm. long, the early whorls are cancellate-
nodulose, but on 4th and 5th whorls the nodules become very feeble and even-
tually obsolete, the subsutural costa is only slightly developed, the subsutural
groove is scarcely concave; the major costae (3) on body whorl and 5 or 6 of
those on base are distinct in one specimen, feeble in another, and scarcely
stronger than lirae in the other two.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 169
The Dassen Island specimen (S. Afr. Mus. no. 6886), 28 mm. long, 4
whorls, aperture plicate, has the whorls distinctly nodulose but the nodules
peter out on later part of 4th whorl; and each one of the major costae tends
to be divided by an incised stria. This latter feature is very noticeable in the
Lambert’s Bay specimen (S. Afr. Mus. no. 9686).
In two juveniles, 12 and 19 mm. long, from Mossel Bay, only the 1st, 2nd
and part of the 3rd whorl are cancellate-nodulose; subsutural costa not
developed, major costae merely indicated. Two other slightly larger specimens
from the same locality are typical tigrina in sculpture and coloration.
In the ‘tigrine’ form there are on the 5th and 6th whorls usually only 2
costae visible below the subsutural groove; but sometimes the succeeding whorl
may recede far enough to expose, partly or wholly, a 3rd costa.
‘Cominella’ angusta Sow.
1886. Sowerby. 7. Conch., v, p. 4.
mon? id.)\ Mar, Sh. S. Ajr., p. 10, pl. 1, fig.8.
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 49.
1932. Turton. Mar. Sh. Pt. Alfred, p. 53.
Aperture shorter than spire. Protoconch ? Postnatal whorls ? (Sowerby:
8 incl. protoconch). Sculpture clathrate; axial riblets 11 on 1st whorl, increasing
to 15 on 5th [largest specimen seen by me]; crossed by spiral lirae, 4 on Ist
whorl increasing to 6—7 on last whorl, with 8—-g additional ones on base.
Columella curved, no parietal callus; no fasciole: outer lip not indented.
12 X 4°75 mm.; Sowerby: 14 x 4; Turton: 15 mm.
Operculum and radula ?
(When fresh) yellowish with reddish spots: ‘in a single or double row just
below middle of whorl’ (Sowerby); one ‘on every alternate rib just above the
suture’ (Turton); (when worn) pure white.
Dead: Port Elizabeth (Sowerby); Port Alfred (Bartsch, Turton).
Remarks. S. Afr. Mus. has 2 worn specimens (locality ?) identified by
J. H. Ponsonby.
It is very unlikely that this species will prove to be either Afrocominella or
Burnupena. There is no trace of a lip sinus, but the axial ribs and spiral lirae
appear to number the same as in Mangilia ponsonby: Sow.; the columella,
however, is more curved in angusta, and without any trace of the 2 little pleats.
Gen. EutrHria Gray
1850. Gray. Figs. Moll. Anim., iv, p. 67.
1917. Cooke. Proc. Mal. Soc., xii, p. 232 (radulae).
1918. Iredale. ibid., xiii, p. 34 (radula)
1929. Thiele. Handbuch, i, p. 312.
Remarks. The only South African species whose radula is known is
queketti; but the figure published by Cooke is incorrect, being based on a
mislabelled slide in the Gwatkin collection.
170 ANNALS OF THE SOUTH AFRICAN MUSEUM
Several South African species have been referred to this genus, but have
been transferred to other genera, or even families: capensis with syn. lacertina
and szmonianus, ? magellani and turton to Afrocominella; eburnea, fuscotincta,
ordinaria to Peristernia (Fasciolariidae); pura to Pyrene (Pyrenidae-Columbellidae) ;
wahlbergi to ‘Purpura’ (Muricidae) ; and clathrata, fallax, and formosa are juveniles
placed in Euthria by Thiele (1925) with a query.
Key to the species
A. Whorls turreted, with a definite nodulose shoulder.
1. Shoulder below middle of whorl. Protoconch diam. 1-1-2 mm. Ist
whorl with 12 ribs, increasing to 18 nodules on last whorl. . . . ponsonbyi
2. Shoulder at middle of whorl. Protoconch diam. 1:5 mm. 1st whorl
with 16-17 ribs, decreasing to 11-13 nodules on last whorl . . . . queketti
B. ‘Profile of whorls straight, without shoulder. . . . . . « .)) gegen
Euthria ponsonbyi Sow.
Fig. 33
1889. Sowerby. 7. Conch., vi, p. 149, pl. 3, fig. 3.
1892. id. Mar. Sh. S. Afr., p. 4, pl. 1, fig. 12.
Protoconch 24 whorls, diam. 1-1-2, alt. 0-8-1 mm. (relatively higher
than in queketti), smooth. Postnatal whorls 7; 1st whorl with 12 ribs, often
obscure, increasing regularly to 16 on 6th and 18 on last whorl, on the later
whorls ceasing to be ribs and becoming merely nodules around the shoulder;
crossed by fine spiral lirae on early whorls, traceable on later whorls between
suture and the shoulder nodules, but becoming obliterated by the growth-
lines; suture close under nodules, i.e. shoulder below middle of whorl, profile
above shoulder concave; on base in juv. 7-8 feeble lirae with fine intermediaries,
in older examples only a few barely traceable (Sowerby: ‘subobsolete’) grooves.
Parietal callus obscurely dentiform, canal as long as rest of aperture, narrow,
with feeble fold at base, recurved, outer lip internally smooth (Sowerby:
‘subobsolete liratum’). 44 X 20 mm. (protoconch and outer lip broken, canal
strongly reflexed so that its tip is not the most anterior part of the shell).
Operculum and radula unknown.
White, irregularly suffused with brown and with brown flames between
the nodules.
Port Elizabeth (Sowerby); Port Alfred (Bartsch, Turton). St. Sebastian
Bay, 27 fathoms, one juv.; off East London, 34 fathoms, one; off Umhlangakulu
River (Natal), 50 fathoms, one (all dead) (S. Afr. Mus. P.F. coll.).
Remarks. The details of the sculpture on the early whorls distinguish this
species from queketti, together with other characters. In Sowerby’s figure the
concavity of the profile above the shoulder nodules seems slightly exaggerated.
The type was a worn specimen with incomplete canal.
The largest P.F. specimen was obtained in Natal waters; the juvenile,
18 mm. long, in St. Sebastian Bay.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA I7I
Euthria queketti Smith
Figsson@)i(9)2.33
rear. omith. 7: Conch., x, p. 1x0, pl. 1, fig. 1.
Not 1917. Cooke. loc. cit., p. 234, fig. 7 (radula, err. = Afrocominella).
Protoconch 24 whorls, diam. 1°5-1:75, alt. 1-1:25 mm. (relatively lower
and broader than in ponsenbyi), smooth. Postnatal whorls 7; 1st whorl with
16-17 ribs, decreasing to 12 (11-13) on last
whorl, but from about 3rd whorl onwards fee >
ceasing to be ribs, becoming nodules around gees WE
: . : : FERRET
the shoulder, increasing in prominence on later ages
whorls; crossed by spiral lirae 4 on Ist whorl
producing a clathrate sculpture, on 2nd only 2
lirae but numerous fine striae, which become
very fine on 4th or 5th whorl and visible only
above the shoulder, finally petering out,
obscured by the growth-lines; base smooth,
with growth-lines but only very faint indications
of spiral lirae; shoulder at middle of whorl,
profile above concave, below straight or slightly
convex. Parietal callus dentiform, canal longer
than rest of aperture, narrow, with nodule or
angular ridge at its base, recurved, outer lip
internally smooth. 55 X 24 mm. Smallest
specimen seen 17°5 X 8-5 mm.
Operculum broadly oval, nucleus apical,
incurved, 14 X 8 mm. in 55 mm. shell.
Creamy-white, irregularly suffused with Fic. 33.
brown, and with brown flames, shoulder nodules Eythria queketti Smith, juvenile
pale, operculum deep amber-brown. 17°5 mm., with protoconch of
‘ . £. ponsonbyi Sow. for comparison.
Radula with c. 160 rows, central plate tri- ee y
angular, length a little less than basal width, with 3 cusps, the middle one
longer than its neighbours, lateralplate with outer cusp larger than the inner.
Off Durban, 40 fathoms [from fish stomachs] (Smith; also $. Afr. Mus.
Ross-Frames coll.).
Off Umhlangakulu River (Natal), 50 fathoms, 1 dead, 1 living; off
Umbhloti River (Natal), 40 fathoms, 1 juv.; off Cone Point and off O’Neil Peak
(Zululand), 34 and 55 fathoms, 1 each, dead; Algoa Bay, 25 fathoms, 1 living
(S. Afr. Mus. P.F. coll.).
Remarks. The size and shape of the protoconch and the sculpture of the
early whorls are important features distinguising this species from ponsonbyt.
Also, in queketti the ribs decrease in number, whereas in ponsonby: they increase.
Cooke’s figure of a radula ascribed to queketti is due to some mistake; it is
172 ANNALS OF THE SOUTH AFRICAN MUSEUM
certainly that of an Afrocominella. The true radula is very like that figured by
Cooke for E. cornea, but with the central plate more like that figured for linea,
and the lateral plate with a slightly larger outer cusp.
It is curious and suggestive that Thiele (loc. cit., p. 312) gave the distri-
bution of cornea as ‘Mittelmeer bis Siidafrika’; I am not aware that cornea has
ever been recorded from South Africa. Compared with a shell of cornea in
S. Afr. Mus. (ident. Tomlin) queketti is broader and less fusiform owing to the
stronger nodulose shoulder, and has a longer canal; but Smith (loc. cit., p. 111)
said guekettt was more ‘slender’!
The presence of this species in Algoa Bay is rather unexpected; the P.F.
specimen is the only one, dead or alive, known from west of Natal.
The juvenile, 17°5 < 8:5 mm. with protoconch and 4 whorls (fig. 33),
appears at first sight very different from the adult, because the prominent
nodules around the shoulder are developed only on the last two, or three,
whorls. The size and squatness of the protoconch, and the sculpture of the
early whorls agree exactly with older examples and leave no doubt as to its
identity. There is just a hint on the back of the outer lip of the nodules to come.
Euthria filmerae Sow.
1900. Sowerby. Proc. Mal. Soc., iv, p. 1, pl. 1, fig. 3.
1932. ‘Turton. Mar. Sh. Pt. Alfred, p. 54.
Profile of whorls straight, without shoulder. Protoconch 14 whorls, diam.
not quite 2 mm., smooth. Postnatal whorls 8; 1st whorl with 12-14 axial ribs,
decreasing to 7 on last whorl; crossed by numerous spiral grooves, 4 on Ist
whorl increasing to 11-13 on last whorl, 13-15 additional ones on base.
Parietal callus nodiform, canal rather long, open, oblique, outer lip thin,
internally smooth. 40 X 14 mm. (living); 47 * 16 mm.
Operculum unguiform, nucleus apical, incurved, 13°5 X 4 mm. in
40. mm. shell.
Lower part of body whorl below periphery white, rest of shell brown.
Dead: Pondoland (Sowerby, also S. Afr. Mus.); Port Alfred (Turton).
Off Itongazi River (Port Shepstone area, Natal), 24 fathoms, one living;
off Amatikulu River (Zululand), 25 fathoms, one dead (S. Afr. Mus. P.F. coll.).
Remarks. The living and dead P.F. specimens both have a very thin shiny
periostracum (like a coating of white-of-egg). The former retains the operculum,
but most unfortunately the animal was not preserved.
Fam. PYRENIDAE
1902. Pace. Proc. Mal. Soc., v, pp. 36-154 (Columbellidae, list of species).
1929. Thiele. Handbuch, i, p. 302 (Columbellidae).
1931. Tomlin.” Ann. Natal Mus., vi, p. 436.
Tomlin correctly takes the oldest genus to form the family name.
‘... it is a matter of considerable difficulty to satisfactorily subdivide the
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 173
Columbellidae into genera and subgenera; and the difficulty has been much
increased by the misdirected efforts of the mere conchologist . . . whatever
value a section may have originally possessed, its true significance has in most
cases been entirely lost sight of by subsequent authors, and species have been
scattered about among the various genera and subgenera in an amazingly
haphazard fashion’ (Pace, p. 40). For example, apicata was put into Nitzdella
by Smith, but into Alza by Bartsch.
Thiele admits four genera, each with a distinctive radula. So far as is
known the South Africa species fall into Pyrene and Columbella.
The lateral plate of the Pyrene radula is tricuspid: one cusp about midway
on the plate, varying slightly according to the species (e.g. filmerae, burnupi,
albuginosa, fig. 34), and separated by a varying distance from the two apical
falcate cusps. In the available South African material the only radula of the
Columbella form is that of fulgurans, the lateral plate of which is also tricuspid,
but only the apical one is falcate, the other two being broad, like shark’s teeth.
Species dealt with here, but whose radulae are unknown, are included in
‘Columbella’.
The identification of the smaller species, e.g. Thiele’s species from the
Valdivia collection, is often difficult; and suggested synonymies are provisional.
Pyrene albuginosa (Rve.)
Fig. 34(f)
1859. Reeve. Conch. Icon., sp. 223.
1921. Sowerby. Proc. Mal. Soc., xiv, p. 126, fig. (approximata).
1926. Tomlin. Ann. Natal Mus., v, p. 291, pl. 16, fig. 5 (Mitrella natalensis).
1931. id. ibid., vi, p. 436 (Mitrella approximata).
1932. Turton. Mar. Sh. Pt. Alfred, p. 72 (var. major, nom. preocc.).
1933. id. 7. Conch., xix, p. 370 (nom. nov. var. rietensis).
Protoconch 24-3 whorls (junction with Ist postnatal whorl indistinct),
alt. 0-75, diam. 0-5 mm., smooth. Postnatal whorls 7; no axial or spiral
sculpture, except c. 10-11 lirae on lower half of base and rostrum. Outer lip
thickened, c. 6 plicae within; columella with 4-6 granules. Periostracum thin,
crinkly, scarious. 12 X 4°5 mm.
Pale corneous, uniform, or mottled or reticulated with fawn or orange-
brown, usually an interrupted subsutural band, and 2 spiral series (one peri-
pheral, one infraperipheral) of opaque white spots enclosing between them a
pale non-reticulate band; some specimens uniform with only a pale peripheral
band. Periostracum usually pale, but sometimes amber-brown.
Radula with c. 225 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp rather strong.
False Bay to Algoa Bay, Port Alfred, East London, and Natal (auct. and
S. Afr. Mus.). Natal (natalensis).
Off Cape Vidal (Zululand), 80-100 fathoms, one; off Morewood Cove,
Tongaat, Umhloti, and Umhlanga (Natal), 22-36 fathoms, 14; off Umkomaas
174 ANNALS OF THE SOUTH AFRICAN MUSEUM
River, 40 fathoms, 5 juv.; off East London, 20 fathoms, one; all dead but more
or less fresh (S. Afr. Mus. P.F. coll.). 29° 38'S: 31° E., 49 metres Weta,
Living: False Bay and East London (U.C.T.).
Remarks. The juveniles collected at Still Bay by Dr. Muir clearly show
that natalensis is a synonym.
Specimens in 8S. Afr. Mus. labelled albuginosa and floccata Rve. do not
seem very different, except the latter are larger and broader with less tapering
spire, and are also more strongly marked with orange-brown blotches and
reticulation; both show on the body-whorl 2 spiral series of opaque white spots,
enclosing between them a pale non-reticulated band, also usually a series of
white spots below the suture (on the upper whorls the peripheral spots just
show above the suture of following whorl). Some specimens have more or less
regularly spaced, straight or crinkly, axial bars between suture and periphery;
some are orange-brown with a pale peripheral band, others uniform orange-
brown. Von Martens (1903. D. Tiefsee Exp., vii, pp. 56, 106) mentions uniform
‘scarlet-red’ examples of floccata from Pondoland; and a beach example from
Natal in 8. Afr. Mus. is reticulate with brown, but has the peripheral band
bright pink.
The same mottled and reticulate pattern is found in C. seychellarum von
Martens (loc. cit., p. 105, pl. 5, fig. 17) which, however, is an even more
broadly oval shell than floccata.
7 SING, a
Fic. 34.
Protoconchs of (a) Pyrene kraussii (Sow.); (6) P. burnupi (Smith); (c) P. dianae (Thiele);
(2) ‘Columbella’ hella Thiele. Central and lateral radula plates of (e) P. burnupi (Smith) ;
(f) P. albuginosa (Rve.), lateral plate only; (g) P. filmerae (Sow.), lateral plate only; (h) Colum-
bella fulgurans Lam.
A
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 175
A comparison with apicata Smith is also not irrelevant. The colour-
pattern is very similar, and although the colour is never such a deep brown as
in apicata, living and fresh albuginosa often show a very clear pale brown
mottling. P. albuginosa, however, does not have a bulbous protoconch, and I
have not seen a trace of spiral striae on the last whorl in any specimen.
Pyrene filmerae (Sow.)
Fig. 34(g)
1892. Sowerby. Mar. Sh. S. Afr., p. 21 (sagena, non Rve.).
mgopsuid.. Proc. Mal. Soc., iv, p. 3, pl. 1, fig. 8.
4
Protoconch 2 whorls, alt. and diam. o-8—o-g mm., smooth, slightly lop-
sided. Periostracum forming a crinkled and scarious band below the suture,
giving a somewhat coronate or turreted appearance to the spire.
The characteristic (as Sowerby said; but see also splendidula Sow.) dark
brown band against (below) the suture is usually interrupted by white spots,
which may be so large as to disrupt the band into a series of alternating brown
and white areas; this is particularly noticeable in the Natal shells, some of
which also show triangular brown marks on a white ground instead of the
more usual white spots on a brown ground. The dark sutural band is not very
obvious in the fresh specimen from Durnford Point, which is cream with, on
the body-whorl, a peripheral series of orange-brown spots through which runs a
continuous thin white line, a less distinct series of spots on middle of base,
protoconch pinkish.
Aperture in living examples violaceous. Periostracum dull brown,
obscuring the bright pattern seen in beach examples; in the Durnford Point
example pale amber.
Radula with c. 200 rows, central plate very delicate, lateral plate apically
bifalcate, the proximal cusp small.
Port Elizabeth and Pondoland (Sowerby). Port St. Johns, and Natal
(between Durban and Port Shepstone) (S. Afr. Mus.).
Off Durnford Point (Zululand), 13 fathoms, 3 dead, but one with perio-
stracum (S. Afr. Mus. P.F. coll.).
Living: Umgazana (south of Port St. Johns), littoral (U.C.T.).
Remarks. ‘Turton obtained no examples at Port Alfred, and consequently one
suspects that the Bairstow and Filmer shells originally came from farther north.
The scarious band of the periostracum is seen in the Umgazana shell, but
is more strongly developed in the fresh specimen from Durnford Point. The
latter was identified by Sowerby (3rd) as ‘probably splendidula’ (see Sowerby:
1847. Thes. Conch., i, Columbella, pl. 37, figs. 65, 66). If he had seen the other
two shells from the same haul, which are obviously worn filmerae, he might not
have suggested the Philippine splendidula.
Some specimens are very similar in pattern to the illustration of tringa
Sow. (loc. cit., pl. 37, fig. 62).
176 ANNALS OF THE SOUTH AFRICAN MUSEUM
Pyrene pura (von Martens)
Fig. 35(a)
1903. Von Martens. D. Tiefsee Exp., vii, p. 25, pl. 2, fig. 14 (Euthria p.).
1925. Thiele. ibid., xvii, p. 173, pl. 30 (18), fig. 21 (Columbella helena).
1925. id. ibid., p. 180 (Euthria p.).
Protoconch 2 whorls, alt. and diam. 1 mm., smooth (but all specimens
worn). Postnatal whorls 5; spire subtending an angle of 35°, profile of whorls
gently convex. Growth-lines but no axial sculpture. Fine spiral striae over
the greater part of whorl, but (in the present more or less corroded specimens)
variable, when traceable c. 8 on 3rd whorl, increasing to ¢. 12 on 5th; additional
striae on base ¢. 20 (scarcely traceable on rostrum). Outer lip thickened.
Periostracum thin. 14 X 6 mm.; 17 X 7°5 mm.
Fic. 35.
(a) Pyrene pura (von Martens). (5) P. parhelena n. sp. (c) ‘Columbella’ polyarosus n. sp., apex.
(d) ‘Columbella’ confertilirata n. sp., apex and operculum.
White; two shells with very faint indications of subsutural brown spots
and even fainter peripheral marks. Periostracum pale buff or yellowish.
Radula with c. 200 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp moderately strong, separated rather widely from the
apical cusps.
34° 31'S. 23° 2’ E., 500 metres (von Martens: one); 35° 16’ S. 22° 26’ E.,
155 metres (Thiele: one pura, 2 helena).
Cape Point NE. 4 N. 18 miles, 135 fathoms, one living; Vasco da Gama
Peak N. 71° E. 18 miles, 230 fathoms, one dead; Lion’s Head N. 67° E. 25 miles,
131-136 fathoms, 2 dead; 10 dead without precise locality (S. Afr. Mus. P.F.
coll.).
Remarks. These shells are clearly referable to yon Martens’s species.
Thiele separated as helena two out of three shells taken in the same locality,
seemingly because these two showed spiral striae only on the lower part of the
whorl. His figure of helena resembles that of pura in shape.
1l
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA ey ig
Pyrene parhelena n. sp.
Fig. 35(6)
Protoconch 2 whorls, alt. 1:25, diam. 1 mm., smooth, junction with Ist
postnatal whorl distinct. Postnatal whorls 5; spire subtending an angle of 30°,
profile of whorls nearly straight; growth-lines but no axial sculpture. Extremely
fine spiral striae over whole whorl, from 3rd whorl stronger striae appear on
lower half of each whorl, 4—5 on 3rd whorl, increasing to 6—7 on 5th; additional
striae on base c. 20. Outer lip thickened submarginally. Periostracum thin.
14 X 5 mm.
Pale fawn with faint white spots; periostracum pale.
Radula with c. 200 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp moderately strong, well separated from apical cusps.
Cape St. Francis NE. x E. 4 E. 36 miles, 70 fathoms, one dead; Cape St.
Blaize N. x E. 73 miles, 125 Se one living, 3 eee (S. Afr. Mus. A886,
moe70 (lype). P.F. coll.).
Remarks. Similar in shape to barbara ‘Thiele 1925, but with different
sculpture; sculpture similar to that of helena, but shape different.
Pyrene kraussiu (Sow.)
Fig. 34(@)
1844. Sowerby. Proc. Kool. Soc. Lond., p. 53 (Columbella k.).
1847. id. Thes. Conch., i, Columbella, sp. 99, p. 144, pl. 40, figs. 180, 181.
1848. Krauss. Sidafrik. Moll., p. 109, pl. 6, fig. 11 (Mangelia fulgurans).
1848. id. ibid., p. 122, pl. 6, fig. 17 (cereale Menke in litt.).
1860. Gould. Proc. Bost. Soc. Nat. Hist., vii, p. 334 (fulminea).
1894. Sowerby. 7. Conch., vii, p. 7 (kitchingz).
1897. id. Append. Mar. Sh. S. Afr., p. 10, pl 6, fig. 3 (kitching?).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 56, pl. 37, fig. 5 (alfredensis).
1931. Tomlin. Ann. Natal Mus., vi, p. 436.
1932. Turton. Mar. Sh. Pt. Alfred, p. 70, pl. 17, no. 500 (var. albanyana).
1932. id. ibid., p. 70, pl. 17, no. 504 (helena, non Thiele).
Protoconch 24-3 whorls, alt. and diam. 0-5 mm., smooth, last whorl feebly
keeled, with minute spiral striae below the keel (seen only in unworn examples),
the keel runs down obliquely at junction with Ist postnatal whorl. Postnatal
whorls 4-5; axial ribs 10 on each whorl.
Radula with 110-150 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp rather strong (cf. albuginosa). (krausswu and kitchings
forms examined.)
Table Bay, and False Bay to Durban and Tongaat (S. Afr. Mus.). Off
East London, 32 fathoms (S. Afr. Mus. P.F. coll.).
Living: Port Nolloth, Langebaan (Saldanha Bay), False Bay, and Knysna
(GEE. 'E.).
Remarks. Sowerby (1892, p. 4) doubted whether fulgurans was a Pleuro-
tomid, and Turton (1932) listed it definitely as a Columbella, probably after con-
178 ANNALS OF THE SOUTH AFRICAN MUSEUM
sultation with Tomlin. I have seen examples from Knysna (Krauss’s type
locality), including plump (kraussi) and slender (fulgurans) forms.
Turton’s name helena is preoccupied by Thiele, but the two are quite
different species (cf. pura). ‘Turton said his helena had no zigzag lines, but his
photograph shows them.
Although the characteristic zigzag lines are usually present and easily
visible, in some specimens they are obscured by a uniform chestnut-brown
coloration, with sometimes a series of pale spots below the suture and another
below the periphery (kztchingz).
The Durban specimens, taken alive by Burnup, are more delicate and
translucent than specimens from other parts of the coast, especially the uniform
brown kztching: form from False Bay and the west coast.
aberr. zo Bartsch
1915. Bartsch; loe.1cit:; jo. 57,137. SA:
1931. Tomlin. loc. cit., p. 436 (as kraussi aberr.).
Tomlin regarded 20 as an abnormal kraussw. It is more slender even than
the fulgurans form. Although I have seen no similar aberration among the
numerous examples from False Bay and Still Bay, there is one specimen taken
together with two kraussi at ‘Tongaat which agrees with Bartsch’s description
and figure.
Protoconch 2 whorls (but worn), diam. 0-5 mm., smooth. Postnatal
whorls 5; 1st whorl worn, 2nd and 3rd each with to axial ribs, but evanescent
towards end of 3rd whorl, protractive as in kraussu (Bartsch said retractive,
but see his figure), obsolete on 4th and 5th whorls. 7 x 2°5 mm.
Translucent, with orange-brown zigzag axial lines.
Port Alfred (Bartsch; one specimen; Turton: ‘rare’). Tongaat (Natal),
one specimen (8S. Afr. Mus.).
Pyrene burnupi (Smith)
Fig. 34(d), (e)
1901. Smith. J. Conch., x, p. 112, pl. 1, fig. 2 (Columbella b.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 67, pl. 16, no. 488 (Columbella kowiensis).
Protoconch 3 whorls, alt. 0-6, diam. 0-5 mm., smooth, last whorl with
feeble peripheral keel which runs down obliquely at junction with Ist postnatal
whorl. Postnatal whorls 3-4; axial ribs 12-13 on 1st whorl, 13-14 (15) on
end—4th; crossed by spiral lirae 4 on 1st whorl, 5 on 2nd, 6-7 on 3rd and 7-8
on 4th, additional lirae on base 9-10; intersections forming rounded granules
or beads. 4°5 X 1°5-1°75 mm.
Translucent yellowish, 3-4 red-brown interrupted lines (on the spiral
lirae) around middle of whorl, lower part of base also with dark lines or spots.
Radula with c. 200 rows, central plate delicate, lateral plate apically
bifalcate, proximal cusp well separated from the apical cusp.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 179
Natal (Smith); Port Alfred (Turton).
Living: Durban and Scottburgh (S. Afr. Mus. coll. Burnup).
Pyrene langleyt (Sow.)
1897. Sowerby. Append. Mar. Sh. S. Afr., p. 10, pl. 8, figs. 8, 9 (Columbella I.).
Protoconch 2 whorls, alt. 0-6, diam. 0:5 mm., smooth. Postnatal whorls
4. (natalensis: 5); growth-lines but no axial ribs, occasionally a thickened
growth-line simulates a rib; no spiral sculpture except some (c¢. 10) obscure
striae on base. 4°5 X 2 mm.
Corneous-brown, uniform or with 2 series of white spots, one infrasutural,
one from top of aperture.
Radula with c. 200 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp strong, well separated from apical cusps (cf. burnupr).
Port Elizabeth (Sowerby); Port Alfred (Turton); Kalk Bay and Buffels
Bay (False Bay) (S. Afr. Mus.).
Living: False Bay (U.C.T.).
Pyrene lightfoott (Smith)
1901. Smith. 7. Conch., x, p. 112, pl. 1, fig. 3 (Columbella l.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 67, pl. 16, no. 487 (var. assimilans).
Protoconch 2 whorls, alt. and diam. 0:8 mm., smooth, junction with Ist
postnatal whorl distinct. Postnatal whorls 3. Axial ribs, when present, c. 15
on Ist whorl, ¢. 17 on 2nd, ¢. 17-19 on 3rd; crossed by spiral lirae 7 on Ist
whorl, 8 on 2nd, 9-10 on 3rd, additional lirae on base 18-20; lirae flattened,
broader than the sulci. 7 <x 3 mm; plump example 6 x 3, slender
6-5 X 2-5 mm.
Living: yellowish, each lira castaneous brown, including those on base,
seldom interrupted, a series of pale subsutural spots. Beach examples: pale
buff, 5-7 orange-brown interrupted lines (on the lirae), the interruptions
occurring so as to delimit a series of oblong brown patches; base also with
brown lines.
Radula with c. 150 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp strong, separated from the apical cusps but not so
distant as in burnupi.
Kalk Bay (False Bay) (Smith, and S. Afr. Mus.); Port Alfred (Turton).
Off East London, 22 fathoms, 3 dead (S. Afr. Mus. P.F. coll.).
Living: Algoa Bay, 60 fathoms (U.CG.T.).
Remarks. If the condition of the type (types) in British Museum was no
better than that of the cotypes in S. Afr. Mus., it is not surprising that Smith’s
description did not mention the axial ribs. They are, however, very low and
rounded, and though usually developed on 1st and 2nd whorls, are frequently
obsolete on the 3rd.
180 ANNALS OF THE SOUTH AFRICAN MUSEUM
R. M. Lightfoot of the South African Museum found several dead speci~
mens at Kalk Bay, but the species has not been found living in False Bay by
U.C.T. There are no specimens in the Muir collection from Still Bay.
Pyrene atrata (Gould)
1903. Smith. Proc. Mal. Soc., v, p. 374 (Columbella a.).
1910. Schwarz. Tr. Geol. Soc. S. Afr., Xii, p. 115.
Radula (Durban specimen) with c. 150 rows, central plate very delicate,
lateral plate apically bifalcate, proximal cusp strong, well separated from the
apical cusps.
Fossil: Pleistocene, Port Elizabeth (Schwarz).
Living: Durban (Smith). Morrumbene estuary, Inhambane, and Maxixe,
Portuguese East Africa (U.Q.T.).
One of the Inhambane specimens, 3:3 X 1:75 mm., resembles very closely
the figure of padangensis Thiele (1925. D. Tiefsee Exp. xvii, p. 327, pl. 31 (19),
fig. 19) both in form and coloration.
P. atrata occurs in several forms in the Indo-Pacific region.
Pyrene dianae (Thiele)
Fig. 34(¢)
1925. Thiele. D. Tiefsee Exp., xvii, p. 176, pl. 31 (19), fig. 13 (Columbella d.).
Protoconch 14 whorls, alt. and diam. 0-75 mm., smooth, junction with
Ist postnatal whorl distinct. Postnatal whorls 3-34; axial ribs on 2nd whorl
¢. 22-24, on 3rd c. 28-30; spiral lirae 6 on ist whorl, 8 on 2nd, 10 on grd;
additional lirae on base c. 18. 5 X 2 mm.; Thiele: 4-5 x 1-8 mm.
Radula with c. 200 rows, central plate very delicate, lateral plate apically
bifalcate, proximal cusp strong, well separated from the apical cusps.
84-51 9. 19 37 E., Go metres, one (ihiele):
Off Cape St. Blaize, 125 fathoms, one (S. Afr. Mus. P.F. coll.).
Living: 34° 18’ S. 18° 30’ E. (False Bay), 51 metres (U.C.T.).
Columbella fulgurans Lam.
Fig. 34(h)
1822. Lamarck. Anim. sans Vert., vil, p. 296.
1859. Chenu. Man. Conchyl., i, fig. 1076.
Radula with 130-140 rows, central plate wide, arcuate, lateral plate
tricuspid, the proximal cusp the largest, the apical one falcate.
Living: Mozambique Island (U.W.).
‘Columbella’ pyramidalis Sow.
1894. Sowerby. 7. Conch., vii, p. 370.
1897. id. Append. Mar. Sh. S. Afr., p. 10, pl. 6, fig. 4.
1904. Smith. 7. Malac., xi, p. 22 (adjacens n. sp., listed, sine descr.).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 181
1931. Tomlin. Ann. Natal Mus., vi, p. 437.
1932. Turton. Mar. Sh. Pt. Alfred, pp. 70, 71 (pyramidalis and var. fusca).
1932. id. ibid., p. 71, pl. 17, no. 509 (adjacens), and pl. 17, no. 510 (distincta).
Port Elizabeth and Port Alfred. |
Off Cove Rock and East London, 22-32 fathoms (S. Afr. Mus. P.F. coll.).
‘Columbella’ eulimoides Turton
1932. Turton. Mar. Sh. Pt. Alfred, p. 71, pl. 17, no. 511.
Two dead specimens (one very worn) appear referable to this species and
confirm its validity. They are more slender than pyramidalis, with a longer
base and rostrum. The larger measures 8 X 2:5 mm., with 5 postnatal whorls.
The nuclear apex (? if this is the actual nucleus) alt. 0-4, diam. 0-6 mm.
Off Keiskamma River, 33 fathoms (S. Afr. Mus. P.F. coll.).
‘Columbella’ apicata Smith
1899. Smith. 7. Conch., ix, p. 247, pl. 5, fig. 2.
1932. Turton. Mar. Sh. Pt. Alfred., p. 71, pl. 17, no. 520 (rufanensis); and p. 73, pl. 17,
no. 526 (arcuata).
Although the general colour pattern is the same as in filmerae, the spire is
longer and the whorls do not widen so rapidly; in filmerae the whorls are much
broader and appear as if telescoped.
Three topotypes (S. Afr. Mus. coll. Burnup) lack the distinctive proto-
conchs; they show on the body-whor! 5-6 fine spiral striae, not mentioned in
Smith’s description, which said only the base was striate. cf. Alcira elegans.
Durban (Smith); Port Alfred (Bartsch, Turton).
Two specimens from Delagoa Bay (U.W.), 9°5 < 3:5 mm. (protoconchs
missing) may be this species. Outer lip thickened, plicate within.
One of these shells has the filmerae colour pattern, but the other has
numerous close-set narrow, straight or slightly crinkly axial stripes, brown on a
yellowish ground, about 30 on the body-whorl (cf. cincinnata von Martens, 1880.
Mauritius & Seychellen, p. 248, pl. 20, fig. 14). Margin of outer lip and canal of
both shells chestnut-brown.
‘Columbella’ mutabilis Turton
Smith: lightfoott var., ined. specimens at Brit. Mus.
1932. Turton. Mar. Sh. Pt. Alfred, p. 68, and vars. multicostata and convexa, pl. 16, nos, 489,
490; 491.
Protoconch 2 whorls, alt. 0-75, diam. 0-5-0-6 mm., smooth, junction with
Ist postnatal whorl distinct. Postnatal whorls 4; axial ribs 11-12 on 1st whorl,
increasing to 13-14 (15) on last whorl; crossed by spiral lirae 5 on 1st whorl,
6 on and, 7 on grd, 7-8 on 4th, additional lirae on base 10-12. 7 X 2°75 mm.
Buff or fulvous.
182 ANNALS OF THE SOUTH AFRICAN MUSEUM
Port Alfred (Turton); False Bay and Still Bay (S. Afr. Mus.).
Off Cove Rock and East London, 27-32 fathoms, 4 dead (S. Afr. Mus.
Pie colly):
Remarks. ‘Turton’s var. convexa does not seem worth maintaining, but if
multicostata is retained it must be renamed (preocc. Blankenh. 1901). This
may be a composite species, but further and better material is required. If
Turton had given an illustration of consanguinea Sow. 1897 (included in Mangilia
by Bartsch and Turton) a comparison with mutabilis might have been possible;
there seems to be some resemblance, as far as one can judge from Sowerby’s
figures.
‘Columbella’ polyarosus n. sp.
Fig. 35(¢)
Protoconch 2 whorls, alt. 1-5, diam. 1-3 mm., smooth, with faint axial
pliculae prior to junction with Ist postnatal whorl. Postnatal whorls (largest
specimen) 54; spire subtending an angle of 30°, profile almost straight in early
whorls, becoming gently convex later. No axial sculpture, but growth-lines
distinct in the spiral sulci, less distinct across the lirae. Spiral lirae 7 on 1st-3rd
whorls, 8 on 4th and 5th, additional lirae on base (of 3rd whorl) c. 20. Lirae
flattened, subequal in width to the sulci. Columella slightly curved. Proto-
conch plus 3 whorls 11 X 3°75 mm., protoconch plus 5 whorls 22 x (approx.)
7mm. (Type.)
Cream or buff, the largest specimen with faint orange-brown axial flames.
Off Cape Vidal (Zululand), 80-100 fathoms, one apex; off O’Neill Peak
(Zululand), go fathoms, one with 3 whorls; off Cape Natal, 54 fathoms, one
22 mm. specimen, but last whorl broken; off Umhloti River, 40 fathoms, one
(3 whorls) and 3 fragments; off Hood Point (East London), 49 fathoms, one
(2 whorls); off Cape St. Blaize, 125 fathoms, fragment of apex; all dead
(S. Afr. Mus. A8875-—78, and A8882. Type A8875. P.F. coll.).
Remarks. Has the appearance of a very large Daphnella sulcata, with larger
protoconch and less deep sutures. The aperture is Columbellid, not like that
of Daphnella.
‘Columbella’ confertilirata n. sp.
Fig. 35(d)
Fusiform. Protoconch 2 whorls, alt. 1, diam. 1:25 mm., smooth, faint
axial pliculae prior to the sigmoid junction with 1st postnatal whorl. Postnatal
whorls 4, spire subtending an angle of ¢. 35°, profile of whorls gently convex.
Growth-lines but no axial sculpture; spiral lirae 5 on 1st whorl, 7 on 2nd and
grd, 8 on 4th, additional lirae on base c. 14; lirae rounded, wider than the
sulci. Columella nearly straight. 11 & 4:5 mm.
Operculum narrow oyate, nucleus apical, 2°75 * I mm.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 183
Cream or buff, with very faint indications of orange marks; operculum
dark brown.
Off Umkomaas River (Natal), 40 fathoms, 2, 2 juv., and fragments
foie. Mus: AS88709. P.F. coll.).
Remarks. The shape is similar to that of Mitromorpha volva. ‘The two larger
specimens have 4 whorls, but some of the fragments indicate that 5-whorled
examples with a width of 6 mm. occur.
‘Columbella’ adela Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 174, pl. 30 (18), fig. 24.
Four white specimens from Still Bay (Muir coll.) agree with Thiele’s
description and figure.
34° 51 S. 19 37 E. 80 metres (Thiele). Thiele recorded it also from
Great Fish Bay, Angola.
It may be compared with kincaidi Tomlin 1926, which is yellowish, and
amphitrite ‘Turton 1932, which is brown.
‘Columbella’ meta Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 175, pl. 31 (19), fig. 3.
1925. id. ibid., p. 175, pl. 31 (19), fig. 12 (veneris).
Two dead specimens may be referable to meta, though not quite so slender:
6 X 2mm. as against 7-5 < 2:4 mm.; and with only 5 spiral lirae on 3rd and
4th whorls compared with 6—7 (if Thiele’s figure is exact in this detail).
oh to 9. 22° 26 H..155 metres (Thiele); 94° 51 S. 19° 97’-E., 80 metres
(Thiele: veneris).
S4q20 9. 25 42 EH: 124 fathoms, one; Cape St. Blaize N. x E. 793 miles,
125 fathoms, one (S. Afr. Mus. A8569 and A888r. P.F. coll.).
C. veneris would seem to be extremely close, if not synonymous.
‘Columbella’ brunnescens Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 175, pl. 31 (19), fig. 2.
Two specimens seem referable to this species, 4°55 x 2 mm. and
5°3 X 2:2 mm.
34°38 S. 24°59 E. 80 metres; and 33° 50'S. 25°48’ E. (depth not
recorded) (Thiele).
Off Cape Recife, 56 fathoms; off Cape St. Blaize, 39 fathoms (S. Afr.
Mus. A8580, A8581. P.F. coll.).
‘Columbella’ hella Thiele
Fig. 34(d)
1925. Thiele. D. Tiefsee Exp., xvii, p. 176, pl. 31 (19), fig. 5.
1932. Turton. Mar. Sh. Pt. Alfred, p. 74, pl. 17, no. 531 (brunescens [sic], non Thiele).
184 ANNALS OF THE SOUTH AFRICAN MUSEUM
Protoconch 14 whorls, alt. and diam. 0-5 mm., smooth, junction with Ist
postnatal whorl distinct. Postnatal whorls 4-44; spiral lirae 6—7 on 1st and 2nd
whorls, 7-8 on 3rd, 8-g (10) on 4th, additional lirae on base ¢c. 15; lirae
flattened, wider than the sulci. 6 <x 2 mm.; Thiele: 6-5 x 2-25 mim:
Buff or fawn, protoconch glossy brown; the Natal specimen is glossy
cream (presumably fresher than the others).
35° 16’ S. 22° 26’ E., 155 metres, one (Thiele). Port Alfred, two (Turton).
Off Illovo River (Natal), 27-30 fathoms, one; off East London, 32 fathoms,
12; off Nieca River (East London area), 43 fathoms, 2; off Keiskamma River,
33 fathoms, one; off Great Fish Point, 51 fathoms, one; 34° 5’S. 25° 55’ E.,
67 fathoms, 6; all dead (S. Afr. Mus. P.F. coll.).
Remarks. Fortunately ‘Turton’s brunescens [sic] seems to be the same as
hella, otherwise 1t would require a new name.
‘Columbella’ vitula n. sp.
Fig. 36(a)
Juv. Protoconch 14 whorls, alt. 0-5, diam. o-6 mm., smooth, a few fine
pliculae prior to the indistinct junction with 1st postnatal whorl. Postnatal
whorls 24; axial ribs ¢. 20-22 on each whorl, slightly protractive, evanescent
on base, tops of ribs forming granules separated by a spiral groove; crossed by
spiral lirae 4 on 1st whorl, 5 on 2nd, more distinct in the intervals between the
ribs, additional lirae c. 12 on base (obscure on rostrum). 3°5 x 2mm. Pale buff.
Off Cove Rock (East London), 80-130 fathoms, one juv. (S. Afr. Mus.
AG8e7. EE. colle): 3
Remarks. Although only a juvenile, the sculpture seems distinct and
recognizable enough to justify a specific name, suggested by proximity to the
Buffalo River at East London.
Fic. 36.
(a) ‘Columbella’ vitula n. sp. (6) ‘Columbella’ sigma n. sp., with protoconch further
enlarged.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 185
Resembles C. chuni Thiele (1925. D. Tiefsee Exp., xvii, p. 176, pl. 31 (19),
fig. 6) from the Zanzibar area, 404-463 metres, in having a subsutural groove
interrupting the ribs, but has more ribs.
Thiele’s description of the radula indicates that chun is a Pyrene; the
present species probably also belongs to this genus.
‘Columbella’ sigma n. sp.
Fig. 36(d)
Protoconch 5 whorls, alt. and diam. 0-75 mm., nucleus smooth, whorls
with close-set pliculae, junction with 1st postnatal whorl strongly sigmoid.
Postnatal whorls 24, profile convex, very slightly shouldered; axial ribs 14 on
1st whorl, 15 on 2nd, 8 on the last half whorl, from suture to suture, evanescent
on base of body-whorl. No spiral sculpture except about 8 lirae on lower part
of base and rostrum (the tip of the rostrum seems to be worn). Columella
slightly angular at midway. 4 x 2 mm.
Pale corneous, an opaque white band around shoulder and a disconnected
series of small white streaks from top of aperture, protoconch fawn.
Off Gove Rock (East London area), 22 fathoms, one (S. Afr. Mus. A88qgo.
PF. coll.).
Remarks. Among the known protoconchs of South African species this
seems quite distinctive. Perhaps not a Columbellid.
Fam. RAPIDAE
1929. Thiele. Handbuch, i, p. 300 (Magilidae).
The family is characterized, as far as is known, by the absence of a radula.
Although Thiele adopted the family name Magilidae in place of Coral-
liophilidae, the oldest genus is Rapa Montfort 1810.
Gen. LATIAxIs Swainson
1935. Tomlin. 7. Conch., xx, pp. 180-3 (list of Recent species).
Rapana fritschi von Martens was included in Latiaxis by Tomlin (1923,
1935). Owing to a mistaken identity I removed it to Tritonalia (Muricidae)
(1957), but now return it to the present family in the genus Coralliophila (v. infra).
Smith considered that his Latiaxis rosaceus (1903) might, conchologically,
equally well be put into Coralliophila, but I leave it where Smith originally
put it.
No further examples of L. tortilis or L. capensis Tomlin 1928 have been
found in the P.F. bottom samples recently examined, but the search was
rewarded by one example of a remarkable species which appears to be new.
186 ANNALS OF THE SOUTH AFRICAN MUSEUM
Latiaxis rosaceus Smith
1892. Sowerby. Mar. Sh. S. Afr., p. 16 (nodosus, non Adams).
1903. Smith. Proc. Mal. Soc., v, p. 376, pl. 15, fig. 16.
1923. Tomlin. 7. Conch., xvii, p. 46 (as syn. of fritschi).
1932. Turton. Mar. Sh. Pt. Alfred, p. 78.
1935. Tomlin. loc. cit., p. 183 (as syn. of fritschi).
Aperture (incl. canal) about 14 times spire. Protoconch 2 whorls, alt. 1,
diam. 1-3 mm., smooth. Postnatal whorls 4, profile angularly shouldered at
(or slightly below) middle of whorl. Axial ribs 10 on Ist and 2nd whorls,
10-9 on 3rd and 4th, sometimes only 8 on 4th, usually prominent; spiral lirae
4 on Ist whorl, 6 on 2nd, increasing to g-10 on 4th, 8-9 additional lirae on
base, with intermediaries; main lirae and intermediaries strongly and closely
squamulose (also the interstices when visible). Rostrum umbilicate and
costate, columella slightly curved, anteriorly with free edge slightly reflexed
over umbilicus. Aperture angularly piriform, posterior margin of outer lip
oblique to preceding whorl. 18 X 13 mm. Smith: 21 X 13 mm.
Operculum and animal unknown.
Grey, aperture pale violaceous. Beach specimens rose-pink, salmon, or
white.
Port Elizabeth, Port Alfred (Smith, Turton); Still Bay and Port Alfred
(S. Afr. Mus.).
Off Durnford Point (Zululand), 13 fathoms, one dead but fresh (S. Afr.
Mus. PF: ccoll?):
Remarks. The above description is from the fresh P.F. specimen measuring
15°55 X 10 mm. This was seen by Tomlin and identified as fritscht. ‘Tomlin
did not regard rosaceus as a distinct species in spite of Smith saying that ‘rubro-
coccinea’ [= fritschi] should not be confused with rosaceus on account of the
difference in shape.
In the Muir collection from Still Bay there are a few specimens whose
squat shape and shouldered whorls at once distinguish them from /riéschi.
Approximately equal-sized shells measure: vosaceus 15:5 X 11 mm. compared
with fritschi 16 * 9 mm. I have seen no intergrading examples, and therefore
maintain Smith’s species.
Latiaxis tortilis H. & A. Adams
Fig. 38(a)
1864. H. & A. Adams. Proc. Zool. Soc. Lond. (for 1863), p. 431.
1882. Sowerby. Thes. Conch., v, p. 424, fig. 1 (not the Type, see Smith, 1906).
1903. id. Mar. Invest. S. Afr., ii, p. 228.
1906. Smith. Ann. Natal Mus., i, p. 39.
1935. Tomlin. loc. cit., p. 183 (= gyratus Hinds 1844).
1942. Yen. Proc. Mal. Soc., xxiv, p. 225.
Spire less than aperture (allowing for the missing protoconch about 1}
times in aperture). Postnatal whorls 5; spire turreted, profile of whorls
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 187
angularly shouldered. Sutures undulate. Axial ribs on 1st whorl g (somewhat
worn), and 2nd and 3rd whorls g, on 4th 10, on 5th 10 and i1 irregular and
interrupted by an injury, broad and rounded, from suture to suture, petering
out on base, shoulder keel in apical view undulate; crossed by a strong peri-
phera] keel forming the shoulder, becoming prominent and somewhat laminar
and upturned on last whorl at intersections with the ribs; spiral lirae above
shoulder keel on 1st and 2nd whorls worn, below shoulder on Ist whorl! 2, on
end 2-3, on grd 4—5 above and 4-5 below, on 4th 6-7 above and 6-7 (8) below,
on 5th 8 above and 8-9 below; 20 additional lirae on base; lirae cn body-whorl
subequal; all lirae closely squamulose. Rostrum costate, with 6-7 squamae.
Umbilicus narrow. Aperture plicate within. 43 xX 25 mm. (28 incl. shoulder
projections). Diameter of broken surface at apex 1-5 mm.
Dirty white, aperture pure white.
Vasco da Gama Peak (Cape Point) S. 75° E., distant 134 miles, 166
fathoms (Sowerby, 1903) (S. Afr. Mus. A4g5o. P.F. coll.).
Remarks. The single specimen obtained by the Pieter Faure is here described
and figured; Sowerby only recorded it.
Sowerby identified it with the Chinese ¢ortilis; it was seen by Tomlin,
who, however, made no comment on it when he described L. capensis (1928.
Ann. S. Afr. Mus., xxv, p. 332).
The specific status of tortilis seems to be not satisfactorily decided. Sowerby
disagreed with Gray in making it a synonym of idoleum Jonas; Smith agreed
with Gray. Tomlin makes both zdoleum and tortilis synonyms of gyratus Hinds
1844; Yen keeps forizlis and gyratus separate.
The Type of forizizs is in the British Museum (Cuming coll.); the original
authors stated it had 6 whorls, but gave no size; Yen said it had 7 whorls and
measured 38-5 X 25:4 mm., adding that gyratus (Type also in B.M.) was a
smaller species. The latter statement seems correct: Hinds’s figure shows 6
whorls, including protoconch, and measures only 19 mm. long (assuming his
figure is natural size). Unfortunately Yen did not figure the two species.
The present specimen with 5 whorls (it is unlikely that another postnatal
whorl as well as the protoconch is missing) is larger than the Type of fortilis.
In the original description the spire was said to equal the aperture; here the
aperture is distinctly longer than the spire.
But the present specimen agrees with fortilis in having axial ribs (‘plicis
undulatis distantibus’) whereas according to Hinds’s description and figure
ribs are completely absent in gyratus. I prefer, therefore, to follow Yen in
recognizing two species, and agree with Sowerby in assigning the present
specimen to fortilis.
Apart from the specific identity of this specimen, considerable interest
attaches to its alleged provenance. Both gyratus and tortilis were recorded from
_ the East (Macassar Straits and China). A locality on the slope of the continental
shelf west of Cape Point is indeed surprising.
188 ANNALS OF THE SOUTH AFRICAN MUSEUM
The labelling of the catches on board the Pieter Faure seems to have been
careful. Some, but very few, anomalies in the recorded localities have come
to light; and these are most likely to have been due to faulty transcriptions of
labels when specimens were sent away to specialists.
In the present instance the P.F. label is not available, but the number
P.F. 2561 is entered in the S. Afr. Mus. Register book. The Pveter Faure log-
book gives for this number the locality as recorded by Sowerby (as above), but
does not refer to any Gastropods. Probably therefore the number attached
to the shell sent to Sowerby was an error; but any suggestion as to what was
the correct number is impossible. The original number might have consisted
of five numerals, and the label got torn. For example, P.F. 12561 refers to a
locality off Gape Natal, 185-200 fathoms, which would be far more credible
as the provenance of an example of tortilis; but even that number refers to
other animals in the haul, not Gastropods.
The recorded locality must, therefore, be accepted provisionally, with the
hope that future trawling will obtain further examples of this species in South
African waters.
Latiaxis kylix n. sp.
Fig. 37
Shell obconic, flat above, spire very short, whorls rapidly expanding.
Protoconch 3 whorls, alt. 0.9, diam. 1 mm., on 1st whorl a very feeble keel
below middle of whorl, continued a little more conspicuously on 2nd whorl,
with an additional very feeble one above, on 3rd whorl the latter obsolete and
the lower keel towards end of whorl overlain by the suture of Ist postnatal
whorl, very faint axial pliculae visible chiefly on 2nd whorl, junction with Ist
postnatal whorl marked by a curved varix. Postnatal whorls 3, 1st forming
with the protoconch the spire, feebly keeled, and with fine axial plicae, the
later portion sloping away from spire and passing into the nearly horizontal
2nd whorl; 3rd whorl horizontal above, flat but slightly curved upwards at
the periphery where there are 12 angular, complanate processes; growth-lines
distinct, shortly squamulose at suture with preceding whorl. Base with obscure
spiral striae, visible chiefly near rostrum, which is costate, with 6 squamae.
Bie. 37.
Latiaxis kylix n. sp.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 189
Umbilicus deep. Columella sigmoid. Aperture triangular, canal narrow,
curved. Alt. 10, diam. maj. 18, min. 12 mm. Pale buff.
Off Cape Natal (Durban), W. x N. distant 64 miles, 54 fathoms, one
dead (S. Afr. Mus. A8850. P.F. coll.).
Remarks. Shape resembling the Greek kylix, a widely open cup.
Only this one specimen was obtained although the Pieier Faure carried out
several dredgings in the same area. It is in fresh, unworn condition.
Coralliophila fritscht (von Martens)
Fig. 38(d)
1874. Von Martens. Jahrb. D. Malak. Ges., i, p. 135, pl. 6, fig. 3 (Rapana f.).
1892. Sowerby. Mar. Sh. S. Afr., p. 16 (Pseudomurex meyendorffi, non Calcara).
1903. Smith. Proc. Mal. Soc., v, p. 377 (rubrococcinea, non M. & S..).
?1910. id. Ann. Natal Mus., ii, p. 194, pl. 7, fig. 7 (fragosa).
1910. Stebbing. Ann. S. Afr. Mus., vi, p. 356 (Murex (Pseudomurex) aédonius, non Watson).
1914. Tomlin & Shackleford. 7. Conch., xiv, pp. 246-7 (Pseudomurex meyendorffi, non Calcara).
1923. Tomlin. 7. Conch., xvii, p. 46 (Latiaxis f., not the synonymy).
1932. Turton. Mar. Sh. Pt. Alfred, p. 76, pl. 18, no. 551 (Tritonalia semidisjuncta, = sub-
scalariform aberr.).
1932. id. ibid., p. 79 (C. fritschi and Pseudomurex meyendorffi, non Calcara).
1935- Tomlin. loc. cit., p. 181 (Latiaxis f., not the synonymy).
Not Barnard. 7. Conch., xxiv, p. 180, 1957 = Tritonalia sperata.
Protoconch 5 whorls, alt. I°5, diam. 1-3 mm., 1st whorl smooth, glossy,
2nd smooth but minutely crimped at upper suture, and with faint trace of an
incipient spiral keel, 3rd—5th whorls bicingulate, crimped at upper suture
indicating incipient axial pliculae, 4th and 5th with numerous axial pliculae,
slightly protractive from upper suture, strongly protractive between the two
keels, then retractive, but scarcely traceable to lower part of whorl where there
is a series of minute beads, the lower keel obscured on last part of last whorl by
Ist postnatal whorl, keels minutely beaded where the pliculae cross them,
junction with Ist postnatal whorl marked by a varix. Postnatal whorls 3,
axial ribs 10 on each whorl, prominent on Ist, and becoming flattened, broader,
and indistinct on 3rd; crossed by spiral lirae 4 on 1st whorl, the first 2 not
prominent, the 3rd strongest and peripheral, 5 at end of whorl due to inter-
polation of a thin intermediary below the peripheral lira, on 3rd whorl 7 lirae,
the 4th peripheral, but at end of whorl 8 lirae, the 5th being peripheral; 5
additional lirae on base of 1st whorl (juv.), 7 in the 3-whorled specimen; all
lirae strongly squamate. Rostrum costate and squamate. Columella nearly
straight, slightly rimate anteriorly, a feeble umbilicus. Canal short. Aperture
piriform, the posterior margin of outer lip horizontal and almost perpendicular
to the preceding whorl. One-whorled 5-3 x 3 mm., 2-whorled 7 x 4:3 mm.,
3-whorled 11:5 X 7°5 mm., 4-whorled (minus protoconch) 14 X 9 mm.;
beach-worn 27 X 16mm. Von Martens, ‘fere 6’ whorls 32 x 20 mm. Smith,
fragosa, 6 whorls 28 xX 14 mm.
Operculum oval-reniform, nucleus on outer margin a little below middle.
TGO ANNALS OF THE SOUTH AFRICAN MUSEUM
Creamy-white, operculum amber. Beach shells pink or white.
Only one animal available: no radula was found.
False Bay (von Martens); Port Elizabeth, Port Alfred, East London
(auct.). Still Bay and Tongaat (north of Durban) (S. Afr. Mus.). Off Scott-
burgh (Natal), 168 metres (Stebbing). Scottburgh (Smith: fragosa).
(ey y Ba
mathe oh
Fic. 38.
(a) Latiaxis tortilis H. & A. Adams. (6) Coralliophila fritschi (von Martens).
Off Cape Natal (Durban), 54 fathoms, 1 living, 2 juv.; off Tugela River,
65-80 fathoms, 1 juv.; off Umkomaas, 40 fathoms, 3 and 3 juv.; off Sandy
Point (north of Kei River), 51 fathoms, 1 juv.; off Gove Rock (East London),
22 fathoms, 2 juv.; 33° 3S. 27° 57_E., 32 fathoms, 1 juv.;. 34° 5 S225 pees
67 fathoms, 3 juv.; 34° 27'S. 25° 42’ E., 256 fathoms, 1 juv.; all dead except
one (S. Afr. Mus. A8858—A8865. P.F. coll.).
Kosi Bay, and 33° 37'S. 26°56’ E., 46 metres (U.C.T.).
Remarks. Von Martens gave a recognizable description and figure of a
worn shell (which probably explains why he did not say the lirae were
squamose), which can be matched without any ambiguity by numerous beach
shells from Still Bay (Muir coll.) and Port Alfred in S. Afr. Mus. It seems
strange that the name of a Persian Gulf species (rubrococcinea) should ever have
been dragged into the South African fauna-list.
There is a tendency to subscalariformity, exemplified by two (out of 30)
of the Still Bay shells, which measure (apices worn, only 2nd—5th whorls
present) 24 x 12 and 22 x 13 mm. There are g axial ribs traceable on the
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA IQI
last whorl in both specimens. Turton’s Tritonalia semidisjuncia is obviously
another example. Von Martens compared the general shape of /ritschi with
Purpura (Rapana, Coralliophila) scalariformis Lam.
The single shell from Tongaat measures 15 X 9 mm. (4 whorls proto-
conch missing) and has the convex, non-shouldered profile of fritschiz, though
the spire is rather short. It is not a rosaceus.
Stebbing referred a Natal Hermit-crab to the Tristan species Eupagurus
tristanensis, stating that it inhabited the same species of shell: Murex (Pseudo-
murex) aédonius Watson. In general appearance there is certainly much
similarity between aédonius and fritsch: (and fragosa), but the former has fewer
(8) axial ribs. Stebbing did not return the shell with the crab, but I think there
is no doubt that it was an example of fritschr. (The crab was later referred to
an Indo-Pacific species, not the ‘Tristan species.)
I strongly suspect that fragosa is only a slender form of fritschi, with g axial
ribs (as in the above subscalariform examples), less convex profile, and posterior
margin of the outer lip oblique. Smith said the spiral lirae numbered about
14 on the penultimate and about 36 on the body whorl; the figure shows only
6 and 20 respectively, which agrees very nearly with the number in normal
Jritschi; but possibly the artist did not insert all the fine intermediaries.
The shape of the shell and of the aperture indicate that the above described
3-whorled shell and the juveniles are indubitably /fritschi. The details of the
axial ribs and spiral lirae are the same, though in 5-whorled shells the lirae
may increase to 8—-g, with 7-8 additional ones on base, and the anterior part
of the columella is slightly reflexed over the umbilicus.
Among the beach-worn shells from Still Bay (Muir coll.) is one 4-whorled
shell which shows the curved varix at end of the protoconch, preceded by a
faint indication of a spiral keel.
The presence of 13 protoconchs in 8 bottom-samples, and one living
specimen, in the area between Durban and Algoa Bay, shows that this is by no
means a rare species. Beach examples are fairly common within the same area.
The species is probably a rock dweller, and this may explain why the Preter
Faure obtained only one living example, because she avoided the rough ground
and used a fishing-trawl more often than a dredge.
Coralliophila isosceles n. sp.
Fig. 39(2)
Protoconch 14 (2) whorls, alt. 1, diam 1-25 mm., smooth, a faint peri-
pheral keel on last part, junction with 1st postnatal whorl indistinct. Postnatal
whorls 4-44, profile of spire straight, periphery at bottom of whorls, below
which the whorl contracted to the sunken suture. No axial ribs. Spiral lirae
on Ist whorl 3, the 3rd peripheral and strongest, on 2nd whorl 5, the 4th peri-
pheral and strongest; on 3rd 5 or 6, the 4th or 5th strongest; on 4th whorl 9,
the 6th strongest; 8-g additional lirae on base; all lirae squamose. Aperture
192 ANNALS OF THE SOUTH AFRICAN MUSEUM
angularly piriform, contracted anteriorly, rostrum costate, canal open, colu-
mella with free edge anteriorly, a shallow umbilicus. 18 (with protoconch) x
12 mm.; 15 (with protoconch) x 10-5 mm. Pale greyish-brown.
Off Glendower Beacon (Port Alfred area), 66 fathoms, 2 dead but fresh
(S. Afr. Mus. A4g51. P.F. coll.).
Remarks. In general shape somewhat similar to rosaceus, but easily distin-
guished by the straight profile of the spire, the peripheral lira near bottom of
whorl, and the absence of axial ribs or knobs.
Fic. 39.
(a) Coralliophila isoceles n. sp. (b) C. zuluensis n. sp., whole shell to illustrate shape (squamae
on lirae not indicated), and protoconch.
Coralliophila zuluensis n. sp.
Fig. 39(4)
Protoconch 3% whorls, alt. 0-6, diam. 0-75 mm., 2nd and 3rd whorls each
with c. 30 pliculae, cut by 2 spiral striae, the upper one feeble and indistinct,
on last half whorl the lower stria and the pliculae below it are concealed by the
encroaching suture of the Ist postnatal whorl, the protoconch is consequently
lopsided, junction with ist postnatal whorl marked by a slight varix. Postnatal
whorls 6, profile of first 4 carinately angular, of last 2 convex; axial ribs on Ist
whorl 10, on and and grd 11, on 4th 9g, on 5th and 6th 8, broadly rounded,
petering out on upper part of base; crossed by spiral lirae 4 on 1st whorl, the
3rd lira most prominent, on 4th and 5th whorls 5, the 4th most prominent, on
5th and 6th 7, the 5th lira a little more prominent than the others, in places 2
12
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 193
additionai lirae are visible one above and one below the peripheral lira; on
upper whorls the strong peripheral lira forms projecting knobs at the inter-
sections with the axial ribs, on later whorls it merely accentuates the roundness
of the ribs; on base 6 additional lirae; all lirae strongly squamulose. Rostrum
costate and squamose; canal short, distinctly delimited, partly overgrown by
anterior end of columella. 21 x 13 mm.
Uniform cream, interior of cana] mauve.
Off O’Neill Peak (Zululand), 90 fathoms, one dead but fresh (S. Afr.
Mus. A8852. P.F. coll.).
Coralliobta madreporarum (Sow.)
1832. Sowerby. Gen. Moll. Purpura, fig. 2.
1859. Chenu. Man. Conchyl., i, figs. 851, 852.
1880. Von Martens. Mauritius G Seychellen, p. 237 (references).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 173, pl. 7, fig. 11.
One living example, 15 <X g mm., white with violaceous columella, rose-
madder operculum Delagoa Bay (U.W.).
Distribution. Mauritius, Réunion, East Indies.
Fam. MURICIDAE
1929. Thiele. Handbuch, i, p. 287.
Murex brevispina Lam.
Figs. 40(b), 41(a)
1822. Lamarck. Anim. sans Vert., vii, p. 159.
1952. Braga. Anais Est. zool. Ultramar., vii, 3, p. 76, pl. 3, fig. 5.
Protoconch 24 whorls, alt. and diam. 1-3~-1-5 mm., smooth, not sharply
demarcated from Ist postnatal whorl.
Radula with c. 120 rows, central plate with median cusp longer than side
cusps, lateral plate stout (cf. ternispina: 1911. Schepman, Siboga Exp. monogr.,
xlix, pl. 24, fig. 8).
Dead: Durban Bay (Krauss).
Living: Delagoa Bay (K.H.B. coll.; also Braga, and U.W.); Inhambane
poe. T.).
Murex fallax Smith
Fig. 41()
1901. Smith. 7. Conch., x, p. 113, pl. 1, fig. 9.
1903 (July). Sowerby. Mar. Invest. S. Afr., ii, p. 227.
1903 (Oct.). Smith. Proc. Mal. Soc., v, p. 375.
Protoconch large, 24 whorls, alt. and diam. 2 mm., smooth. Postnatal
whorls 6 (Smith), 1st whorl sharply demarcated from protoconch by a varix;
194 ANNALS OF THE SOUTH AFRICAN MUSEUM
axial ribs obscure on ist whorl, c. 14 on 2nd, 11 better developed on 3rd;
varices not prominent, 3 on each of the later whorls, shoulder bluntly nodular,
3 (sometimes only 2) intervening ribs bluntly nodular at shoulder and at lower
end; each varix with only one spine a little above middle of rostrum from 3rd
whorl onwards (very feebly developed on 2nd whorl); spiral lirae more promi-
nent on early whorls than the axial ribs, 4 on 1st whorl, on 2nd and 3rd 4 with
intermediaries (total 6—7), obscure on later whorls, on base and rostrum of
3rd whorl (juv.) ¢. 24, on 5th whorl of large specimen few and feeble, outer
surface of reflexed columella (which forms the inner lip) corrugate; imner
columellar surface not lirate. 78 x 41 mm. (Smith); 59 (5 whorls, protoconch
missing) X 30 mm.; protoconch plus 2 whorls 13 x 7 mm.; protoconch plus
3 whorls: 18 X I0 mm.
Fic. 40.
Central and lateral radula plates of (a) Murex ramosus Linn.; (b) M. brevispina Lam.; (c) Trito-
nalia puncturata (Sow.); (d) Trophon acceptans n. sp.; (e) Drupa squamilirata (Smith); (f) Thais
capensis (Petit); (g) T. dubia (Krss.); (h) T. wahlbergi (Krss.); (i) T. castanea (Kiister); (7) Urosal-
pinx heptagonalis (Rve.).
CONTRIBUTIONS TO KNOWLEDGE OF 58.A. MARINE MOLLUSCA 195
Operculum broadly oval, 14 x 10 mm. in 59 mm. shell, nucleus below
centre, growth-lines prominent.
Buff with obscure brown spiral bands, chiefly on the shoulder knobs, and
on rostrum, aperture white, operculum amber-brown.
Dead: off Durban, 40 fathoms, from fish stomach (Smith) ; off Umtwalumi
River (Natal), 25 fathoms, 2 juv. (S. Afr. Mus. P.F. coll.).
Living: off Port Shepstone (Natal), 36 fathoms (Sowerby).
Fic. 41.
(a) Murex brevispina Lam. (b) M. fallax Smith. (c) M. axicornis Lam. (d), (e) Murex sp., whole
shell, with protoconch and Ist postnatal whorl further enlarged. (f) Murex sp. juv.
Remarks. This species has a remarkably large protoconch for a species of
this genus, distinctly larger than that of brevispina, axicornis, and ramosus. By
this character, and by the sculpture of the apical whorls, juveniles are easily
distinguished from brevispina.
Unfortunately the animal of the specimen recorded by Sowerby was not
preserved.
196 ANNALS OF THE SOUTH AFRICAN MUSEUM
There are some Ceylonese specimens in S. Afr. Mus. labelled haustellum,
with pink apertures (? chrysostoma). None of them has a protoconch; but the
ist whorl is narrower than that of fallax, and presumably the protoconch was
also narrower. Axial ribs 14 on 1st and 2nd whorls, 13-14 on 3rd; spiral lirae
only 3 on ist and 2nd whorls, 4 plus intermediaries on 3rd whorl.
Smith said fallax differed from haustellum in colour, but except the white
aperture the large Natal example resembles the Ceylonese specimens.
Murex virgineus Bolten-Roding
1798. Bolten-Roding. Mus. Bolt., p. 141.
1822. Lamarck. Anim. sans Vert., vii, p. 171 (anguliferus).
1931. Lamy. Bull. Mus. Paris (2), iii, p. 304 (angulifer [sic]).
1952. Satyamurti. Bull. Madras Govt. Mus., n.s. 1, 2, pt. 6, p. 155, pl. 15, figs. 1a, 15 and
var. ponderosus Sow.
Ponta Gea, Beira (Lamy).
Murex ramosus Linn.
Fig. 40(a)
1758. Linne. Syst. Nat., ed. 10, p. 747, no. 448.
1822. Lamarck. Anim. sans Vert., vii, 160 (inflatus, non Brocchi).
1880. Von Martens. Mauritius G Seychellen, p. 231 (inflatus).
1952. Braga. Anais Est. zool. Ultramar., vil, 3, p. 76 (inflatus).
Protoconch 2 whorls, alt. and diam. 1 mm., smooth.
Radula with c. 225 rows, central plate with median cusp larger than side
cusps, lateral plate rather slender.
Dead: Port St. Johns (very worn) (S. Afr. Mus.); off Itongazi River
(between Port Shepstone and Port Edward, Natal), 25 fathoms, one juv. very
worn (S. Afr. Mus. P.F. coll.). Delagoa Bay (Braga; and U.W.).
Living: Inhambane (U.C.T.).
Murex axicorns Lam.
Fig. 41(c)
1822. Lamarck. Anim. sans Vert., vii, p. 163.
1903. Sowerby. Mar. Invest. S. Afr., ii, p. 227.
1906. Smith. Ann. Natal Mus., i, p. 38.
1911. Schepman. Siboga Exp. monogr., xlix, p. 346, pl. 24, fig. 11 (radula).
Protoconch 24 whorls, alt. and diam. 1-5 mm., smooth. Postnatal whorls
6, 1st sharply demarcated from protoconch by the spiral lirae but protoconch
has also 2-4 fine axial plicae before the junction. Axial ribs 11 on 1st whorl,
but indistinct, 10 on 2nd, 9 on 3rd and following whorls, every third rib
becoming a varix, the intervening 2 (sometimes in later whorls only one)
becoming peripheral knobs, not crossing base in later whorls; each varix with
a strong but slender pinnate shoulder spine, followed by 5 (3 on outer lip, 2 on
canal) smaller spines, all grooved and hollowed in front; spiral lirae 5 on Ist
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 197
whorl, on 2nd and following whorls intermediaries develop so that whole whorl
is covered with numerous fine lirae; canal nearly closed. Length 70 mm.
(S. Afr. Mus.); 78 mm. (protoconch missing) (U.C.T.).
Operculum broadly oval, nucleus apical, growth-lines prominent.
Buff, with or without brown blotches, or white.
Radula (number of rows ?) central plate with median cusp broader but
not longer than side cusps, lateral plate slender (Schepman).
Living and dead: off Cape Natal, 47 and 54 fathoms: off Umhloti River,
40 fathoms; off Umvoti River, 56 fathoms (Sowerby, and S. Afr. Mus. P.F.
Gallen 29 30 5. 31 29’ E.., 68 metres (s.s. Africana I).
Remarks. ‘The depth ‘110’ fathoms in Sowerby was a typ. err.; it was
repeated by Smith.
Three specimens in S. Afr. Mus. from the Moluccas vary a little from the
South African specimens: two have 2 spines on the canal, the third has 3, and
all have only one large spine with 3—4 minor ones on the outer lip.
Murex adustus Lam.
1822. Lamarck. Anim. sans Vert., vii, p. 161.
1859. Chenu. Man. Conchyl., i, fig. 578.
1911. Schepman. Siboga Exp. monogr., xlix, p. 346.
1938. Adam and Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. Il, 19, p. 155.
1952. Satyamurti. Bull. Madras Govt. Mus., n.s. I, 2, pt. 6, p. 156, pl. 15, figs. 2 a, b (not
good).
Canal about 14 times as long as aperture. Varices with pinnate and
frondose spines. One prominent rounded knob between each pair of varices.
Up to 70 mm.
Ochraceous or fulvous, more or less suffused with smoky-brown or black,
especially the frondose varices.
Off Tugela River, 14 fathoms, one worn and discoloured, 37 x 22 mm.
fae ics Wins. P.F. coll.).
Ber 4775. 30° 29 E., 24 fathoms (U.G.1.).
Remarks. The U.C.T. specimen is covered with a very thin layer of
Sponge.
A very similar Indo-Pacific species: rubzginosus Rve., which has two knobs
or short axial ribs between each pair of varices and is not so ‘sun-burnt’ in
coloration, occurs on the East African coast at Lamu (S. Afr. Mus. coll.
E. L. Layard on board H.M.S. Castor 1856).
Murex sp.
Fig. 41(d), (e)
Canal twice as long as aperture, rostrum elongate. Protoconch large,
lopsided, 2 whorls, alt. 2-5, diam. 2 mm., smooth, ending with a plain narrow
varix. Postnatal whorls 4; axial ribs 12 on each whorl, every 4th rib forming
198 ANNALS OF THE SOUTH AFRICAN MUSEUM
a sharp varix, with a simple slightly curved hollow spine at shoulder; spiral
lirae 3 on ist whorl, uppermost one forming the shoulder, following whorls
with lirae above shoulder and intermediaries below, on last whorl respectively
8 and 10-12, the shoulder lira forming complanate nodules at intersections
with the ribs between each pair of varices; about 8 additional lirae on base and
rostrum, with intermediaries; edge of varices below shoulder spine serrate-
crenulate. Growth-lines between the varices forming a cancellate-granulate
sculpture. Aperture rimate. Canal nearly closed. 37 < 14 mm.
Operculum broadly oval, 6-5 x 4°75 mm., nucleus near apex, growth-
lines prominent.
Radula with c. 130 rows, central plate with median cusp longer than side
cusps, lateral plate slender.
Off Cape Natal, 85 fathoms, one living (S. Afr. Mus. A8833. P.F. coll.).
Murex sp. juv.
Fig. 41(f)
Protoconch 2 whorls, alt. 1:5, diam. 1-25 mm., smooth, ending in a plain
narrow varix. First postnatal whorl with 3 varices, each with a simple, curved,
hollow shoulder spine, and 3 nodules between each pair. Total length 5 mm.
Off Cape Natal, 85 fathoms, one juv. (S. Afr. Mus. A8834. P.F. coll.).
Although taken in the same haul as the previous species, this juvenile
belongs to a different species because the protoconch is much smaller. Larger
specimens might show some resemblance to the East Indies falcatiformis ‘Thiele
(1925. D. Tiefsee Exp., xvii, p. 168, pl. 30 (18), fig. ro).
‘Murex’ uncinarius Lam.
Fig. 42(a)
1822. Lamarck. Anim. sans Vert., vii, p. 166.
1840. Sowerby. Proc. Zool. Soc. Lond., p. 143, and Conch. Illustr., no. 53, fig. 76 (Murex
capensis).
1848. Krauss. Stidafrik. Moll., p. 112 (capensis ? uncinarius).
1903. Von Martens. D. Tiefsee Exp., vii, p. 24 (Murex (Pteronotus) [sic] u.).
1925. Thiele. ibid., xvii, p. 168 (Murex (Pterymurex) u.).
1929. id. Handbuch, i, p. 299 (Tritonalia (Poropteron) u.).
Protoconch 14-2 whorls, alt. and diam. o-8—o-g mm., smooth. Postnatal
whorls 6, 1st sharply demarcated from protoconch; 1st whorl with 1o axial
ribs, 2nd with 8, but towards end of 2nd the alternate ribs begin to become
carinate varices with hollow, alate-uncinate expansions, 3rd and later whorls
each with 3 varices alternating with 3 low rounded peripheral convexities
(scarcely bosses), profile evenly convex to the convexity (no angular shoulder) ;
on last 3 varices the uppermost (largest) alate expansion more or less uncinately
curved upwards towards apex, exsert (not incurved); 2-4 (ocasionally 5)
smaller acute processes below the large uppermost expansion; on preceding
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 199
whorl these processes (except one at suture) are concealed by the succeeding
whorl; spiral lirae on 1st whorl 2 peripheral, obsolete on 2nd whorl, replaced
on this and following whorls by numerous fine striae, particularly well marked
on hinder side of the varical expansions; on 5th and 6th whorls the peristome
appears to become continuous and the canal closed at the formation of each
successive varix. 27 (6 whorls, protoconch missing) X 12 (excl. processes)
Ig mm. (incl. processes); smallest specimen seen (protoconch plus 3 whorls)
17°5 X 2°5 mm. (excl. processes).
Operculum and animal unknown.
Pure white, porcellanous. Beach specimens tend to become buff, pinkish,
or brown.
Port Elizabeth, Port Alfred, Still Bay, False Bay (auct. and S. Afr. Mus.).
St. Francis Bay, 80-100 metres (von Martens); 35° 29'S. 22°26’ E., 155:
metres (Thiele). Algoa Bay, 25 fathoms; off Cape Morgan, 87 fathoms, off
Cape Natal, 54 fathoms (S. Afr. Mus. P.F. coll.).
Bf 115 0. 255 E., 6 fathoms (U.C.T.).
Remarks. ‘The Pieter Faure took four specimens (including the largest
27 mm.) in Algoa Bay, two of them in fresh condition with protoconchs; two
from off Cape Morgan are also fresh with protoconchs and well-marked spiral
striae; the Natal specimen is complete but encrusted with Serpulids.
The locality Table Bay (S. Afr. Mus.) is not acceptable.
So far as I am aware the radulae of this species and the following mitrae-
jformis are unknown. The correct genus therefore remains uncertain: Thiele in
Fic. 42.
(a) ‘Murex’ uncinarius Lam., two views of protoconch. (5) ‘M.’ mitraeformis Sow. (c) ‘M.’
wahlbergi Krss.
200 ANNALS OF THE SOUTH AFRICAN MUSEUM
1925 put uncinarius in Murex section Pterymurex (Rov. 1899, a synonym of
Pterynotus Swainson 1840), but in 1929 in Poropteron Jouss. 1880, a section of
Tritonalia.
‘Murex’ mitraeformis Sow.
Fig. 42(5)
1841. Sowerby. Proc. Zool. Soc. Lond. and Conch. Illustr., fig. 75 (quoted from Krauss) .*
1848. Krauss. Stidafrik. Moll., p. 112.
1892. Sowerby. Mar. Sh. S. Afr., p. 3 (quotes Conch. Illustr., fig. 75).
In general similar to uncinarius but more turreted owing to the whorls
being tabulately shouldered (with an obscure ridge) and sutures deeper.
Protoconch 14 whorls, alt. and diam. 0:75 mm., smooth. Postnatal
whorls 5, 1st sharply demarcated from protoconch; 1st whorl with 9 axial ribs,
and with 7, 3rd—5th each with 3 varices alternating with 3 bosses. The upper-
most and largest expansion on each varix is more tabulate than alate, and
strongly uncinately curved towards apex, the tips incurved and often touching
the preceding whorl; below the large uppermost expansion on last 3 whorls
are 6 smaller processes, blunt, hook-like and curved forwards, at least 2 of
which (sometimes 3) are exposed; spiral lirae 2 on 1st whorl, 3 on 2nd, becoming
4 on 3rd, and thereafter replaced by numerous spiral striae. Canal closed.
22 (protoconch missing) < 9 mm.
Operculum and animal unknown.
Cape and Natal (Sowerby, Krauss) ; Still Bay, Port St. Johns, and Tongaat
(Natal) (S. Afr. Mus.). The west coast of the Cape Peninsula (S. Afr. Mus.)
is scarcely acceptable as a locality.
Remarks. One of six beach specimens from Still Bay retains the protoconch
and spiral striae.
‘Murex’ wahlbergi Krss.
Fig. 42(c)
1848. Krauss. Siidafrik. Moll., p. 111, pl. 6, fig. 13.
1892. Sowerby. Mar. Sh. S. Afr., p. 3 (Trophon w.).
Protoconch 2 whorls, alt. and diam. 1-3 mm., smooth. Postnatal whorls
6, profile from 2nd or 3rd whorl onwards angularly shouldered, 1st whorl
sharply demarcated from protoconch. Axial ribs on 1st whorl 14, on gnd 13-12,
on 3rd 10-9, on 4th, 5th and 6th 9-8, 6th whorl sometimes with only 7 ribs,
from suture to suture, and extending across base to rostrum; from the shoulder
downwards on 3rd and following whorls the ribs are lamellate, somewhat
variable but when well developed sharply pointed at shoulder, and hollowed
* Krauss’s reference to Sowerby seems to be incorrect, though Sowerby himself quoted
fig. 75. Sherborn does not list ‘mitraeformis Sow.’ except as a Scalaria 1844, only mitraeformis
Brocchi 1814. and mitriformis Wood 1828, nom. nud. Sowerby, Proc. Zool. Soc., 1840, p. 143,
gives ‘fig. 75’ for his cancellatus.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 201
in front; spiral lirae on 1st and 2nd whorls 3, almost as strong as the ribs, with
a feeble one between suture and shoulder on 2nd whorl, on 3rd 4 lirae with
2-3 feeble ones above shoulder, on 4th 4-5 or 6 lirae with 4-5 feeble ones
above shoulder; lirae becoming broad and flat with narrow intervals, so that
on later whorls the sculpture is more correctly described as striate. Sculpture
thus cancellate on first 2 whorls (with transversely oblong hollows), axially
ribbed on later whorls and lirate/striate in the intervals. Rostrum rimate.
Canal when fully developed nearly closed. Outer lip plicate within.
AI X 22 mm.
Operculum and animal unknown.
Rubescent, aperiure pale reddish or violaceous (Krauss). Natal (Krauss).
Remarks. Intermediary striae are developed on the body whorl, producing
a somewhat similar sculpture to that of ‘Purpura’ wahlbergi, but the striae are
never so numerous and close together, nor crispate, as in the latter species;
young worn shells might, however, be difficult to identify.
Four adult shells in S. Afr. Mus. were presented by C. A. Fairbridge (who
lived in Gape Town) in 1887, and were registered as coming from Table Bay.
This locality is very doubtful as no specimens have since been recorded from
any Cape locality.
There are also in S. Afr. Mus. 5 juveniles, 6-5-19 mm. long, without
locality but probably collected in Natal (ex coll. Juritz).
Until the animal is discovered Sowerby’s suggestion to transfer this
species to Trophon is only provisional.
Trophon carduus (Brod.)
1832. Broderip. Proc. Zool. Soc. Lond., p. 175 (Murex c.).
1903. Sowerby. Mar. Invest. S. Afr., 1, p. 227 (Trophon c.).
1906. Smith. Ann. Natal Mus., i, p. 38 (Coralliophila c.).
Protoconch 14 whorls, alt. 0:5, diam. 0-75 mm. (but slightly worn),
smooth. Postnatal whorls 6. Axial ribs 11 on all whorls, with angular shoulder,
continued across base on body whorl; spiral lirae one on 1st whorl (at shoulder),
2 on 2nd (upper one forming the shoulder), 3 on 3rd and following whorls,
intersection of shoulder lira with ribs on 1st and 2nd whorls forming little
points, on 3rd and following whorls forming short blunt spines, grooved and
hollowed in front, 3rd lira with shorter spines, 2nd lira with vaulted scales; on
5th and 6th whorls an intermediate lira between 2nd and 3rd lirae, with
vaulted scales; 6 main lirae, with intermediaries, on base, all scabrous; on 5th
whorl between suture and shoulder 4, and on 6th whorl 5 spiral series of vaulted
scales marking the lines of growth. Columella reflexed, canal open, probably
subequal in length to aperture (tip broken). 23:5 (canal tip broken) x 13 mm.
Operculum broadly oval, nucleus apical, 6 x 4 mm.
White, operculum amber-brown.
Living: off Port Shepstone (Natal), 250 fathoms (Sowerby).
202 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. ‘The above description is from the specimen recorded by Sowerby
(now in S. Afr. Mus.) ; it does not correspond with Broderip’s brief description,
which said ‘sexfariam varicosa-spinosa’, and made no mention of the vaulted
scales. On the Natal specimen the spinose varices are undecimfariam and the
shoulder spines increase evenly in length around the whorls, and cannot be
divided into groups of 6 (sexfariam).
Smith suggested the species (? whether he saw the Natal specimen) could
be included in Coralliophila, but Sowerby’s placing in Trophon seems better.
The animal of the Natal specimen was not preserved.
Sowerby said the species was “very rarely met with’, but gave no other
locality than Broderip’s original one from Peru.
The Australian T. carduelis Watson is a very different shell.
Trophon acceptans n. sp.
Figs. 40(d), 43(8)
Protoconch 14 whorls, alt. 1, diam. 0-75-0o-g mm., smooth. Postnatal
whorls 6, profile of first 3 whorls evenly convex, of later whorls angularly
shouldered slightly above the middle; axial ribs on 1st whorl 10, on 2nd 10-11,
on 3rd 11-12, on 4th and 5th 12-13, on 6th 13-14, arching over the suture at
top, extending across base to rostrum, broad basally but sharply keeled on
body-whorl in juveniles and one adult, but abraded in the others, the intervals
in cross-section V-shaped (not U-shaped), the back slope steeper than the
forward slope; no spiral sculpture but a very slight indication of a shoulder
keel between the ribs on body-whorl of adults, and two below even more
obscure. Canal subequal to aperture, open. Protoconch plus 2 whorls
5 X 2°25 mm., protoconch plus 3 whorls 6-5 x 2-5 mm., adult 21 X 9:5 mm.
(figured specimen 18 X 8 mm.).
Operculum oval, nucleus apical.
Pure white, juveniles pale buff, operculum amber.
Radula with c. 100 rows, central plate with median cusp a little longer
than side cusps, a minute denticle between median and side cusp, lateral plate
blade subequal in length to base.
Cape Point E. ? N., distant 36 miles, 630 fathoms, 5 adults; Cape Point
N. 64° E. 37 miles, 700-800 fathoms, one adult; Gape Point E. « N., 35 miles,
500 fathoms, one adult living; Cape St. Blaize N. x E., 73 miles, 125 fathoms,
4 juv. (one living); 34° 26'S. 25° 42’ E., 124 fathoms, 4 juv.; off Cove Rock
(East London area), 80-100 fathoms, one juv., S. Afr. Mus. A3449, A3473
(Type), A3480, A8633, A8634, A884o. P.F. coll.).
Remarks. Of the five remaining adults (two were sent to Tomlin) all are
more or less abraded; four are matt chalky white, the fifth (figured, A3473)
very little abraded on the body-whorl which is almost glossy, especially within
the aperture. The juveniles are clean and fresh.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 203
One juvenile (off Cape St. Blaize) 8 mm., has 10 axial ribs on 1st-3rd
whorls, but only 9 more widely spaced on the 4th whorl. Its radula corresponds
with that of the type.
In describing declinans (1886. Challenger Rep., xv, p. 168, pl. 10, fig. 10)
from 69 fathoms off Marion Island, Watson was inclined to regard it as
conspecific with the North Atlantic truncatus Strom. (see: Sars, 1878. Moll.
Arct. Norveg., p. 246, pl. 15, fig. 9) and separated it only in deference to the
opinion of Sars and Gwyn-Jeffreys. There is a slight but distinct difference in
the convexity of the profile from apex to aperture between the figures of the
two species.
The present specimens (adults) have stronger shoulders than either trun-
catus or declinans, more like clavatus Sars (loc. cit., pl. 15, fig. 12), but the last
species has the ribs acute at the shoulders; nor are the shoulders so high up as
in clathratus Linn. (Sars, loc. cit., pl. 15, fig. 10). The axial ribs are fewer than
in truncatus and declinans.
The Pieter Faure specimens thus seem worthy of acceptance as a species
distinct from declinans. Possibly connecting forms may exist in the area between
South Africa and Marion Island.
There is a strong resemblance to tenuzrostratus Smith (1899. Ann. Mag. Nat.
Hist. (7), iv, p. 241, and 1901. Jilustr. Zool. Investigator. Moll., pl. 10, figs. 4, 4a)
from the Andaman Islands, 185 fathoms. The body-whorl has 3 scarcely
visible spiral keels as in the Cape specimens, but there are one or two fewer
axial ribs, and the upper whorls are shouldered slightly below the middle.
Trophon ? incertus n. sp.
Fig. 43(a)
Protoconch 2 whorls, corroded. Postnatal whorls 5, profile strongly
shouldered about in middle of whorl, sutures undulate; axial ribs 12 on Ist
whorl, 12-13 on 2nd, 13-14 on 3rd, 14-15 on 4th, 16-17 on 5th, from suture to
suture, slightly curved above shoulder, extending across base; crossed by a
spiral lira at shoulder, and one (Type) or 2 (cotype) below shoulder, about 12
additional lirae on base, with an intermediary between each pair; inter-
sections slightly tubercular, strongest at shoulder. Columella nearly straight.
13-14 X 6mm. Pale buff.
Off Cape Natal (Durban), 440 fathoms, two (S. Afr. Mus. A8843. P.F. coll.).
Remarks. Similar to Trophon ? celebensis Schepman (1913. Stboga Exp.
monogr., xlix, p. 452, pl. 30, fig. 13) except that the latter has 5 spiral lirae on
body-whorl, and 20 axial ribs. The Siboga shell was obtained at 462 metres
in the East Indies.
Schepman remarked that it might possibly prove to be a Pleurotomid.
In the present two shells the ribs above the shoulder and the intervening
growth-lines are slightly concave, but scarcely as strong as in most Pleurotomids
(Surcula) ; there is no minute crinkling at the suture.
204 ANNALS OF THE SOUTH AFRICAN MUSEUM
Trophon sp. juv.
Fig. 43(¢)
Protoconch 14 whorls, low, alt. 0-5, diam. 0-8 mm., smooth, a few very
fine pliculae or growth-lines before the definite axial ribs begin. Postnatal
whorls 3, shouldered, profile above and below shoulder straight; axial ribs 14
on Ist whorl, 17 on 2nd, 20 (21) on grd, at shoulder slightly nodular and con-
nected by a lira, feeble on 2nd, indistinct on 3rd whorl, extending not quite
to suture above and obsolete on base. Columella curved, canal rather long.
5°25 X 245 mm. Cream.
Off Cape Natal (Durban), 440 fathoms, one juv. (S. Afr. Mus. A8841.
P.F2 coll.)
Remarks. ‘The shell is extremely fragile. Although there is no trace of a
lip sinus, this shell might possibly be a Surcula.
Fic. 43.
(a) Trophon incertus n. sp. (b) T. acceptans n. sp., with two views of juvenile. (c) Trophon sp. juv.
Trophon acutispira (Sow.)
Fig. 44(¢), (d)
1921. Sowerby. Proc. Mal. Soc., xiv, p. 125, text-fig. (Cominella a.).
1931. Tomlin. Ann. Natal Mus., vi, p. 429 (Cominella a.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 52 (Cominella a.).
1947. Tomlin. 7. Conch., xxii, p. 271 (Afritrophon a.).
Protoconch 2 whorls, smooth. Postnatal whorls 3. The 11-12 axial ribs
are equally as strong as the two spiral lirae, with nodules at the intersections;
4-5 additional lirae on base. 5 mm. long.
Operculum oval, nucleus on outer margin below middle.
13
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 205
Radula with c. 115 rows, central plate with 3 subequal cusps, postero-
lateral corners of base acute, lateral plate slender.
Port Alfred (Sowerby, Turton).
Living: Lambert’s Bay (U.C.T.).
Remarks. The above description from a single specimen (U.C.T.). If it is
correctly identified, acutispira with its smooth protoconch cannot be included
in the genus Afritrophon (v. infra).
Sowerby described two specimens, 10 X 4 mm., with 2 smooth apical
whorls followed by 4 bilirate whorls, base with 4 additional lirae, and axial
ribs forming a ‘crisply nodulous’ cancellate sculpture. Above the shoulder
the whorls are ‘concavely depressed’, a character which apparently induced
Sowerby to place the species in Cominella.
Fic. 44.
(a) Trophon mioplectos n. sp. (b) T. johannthielei n. sp. (c) T. acutispira (Sow.). (f) T. pistillum
n. sp. Central and lateral radula plates of (d) T. acutispira (Sow.); (e) Afritrophon kowieensis
(Sow.).
Trophon jucundus Thiele
1925. Thiele. D. Tiefsee Exp., xvii, p. 169, pl. 30 (18), fig. 13.
1947. Tomlin. 7. Conch., xxii, p. 271 (Afritrophon j.).
Protoconch 14 whorls, alt. 0-75, diam. 0-5 mm., smooth. Postnatal whorls
34, rather squarely shouldered immediately below the sutures; 1st whorl
sharply demarcated from protoconch, with 11 axial plicae, 2nd whorl with 14,
grd with 18 (counting from outer lip upwards the body-whorl has 21); plicae
thin, with free edges, from suture to suture, extending across base to canal; on
end and 3rd whorls 3 spiral lirae, on last half-whorl 4 (by interpolation between
ist and 2nd series); lirae not so prominent as the axial plicae which form
vaulted squamulae at the intersections; 2 additional lirae on upper part of
base, but none on lower part, the plicae on base not or only feebly crinkled. Outer
lip patulate. 5 <x 2-3 mm. (Thiele); 5 < 2-5 mm. (S. Afr. Mus.). Pale buff.
35° 16'S. 22° 26’ E., 155 metres, one (Thiele).
Off Cape St. Blaize (N. x E., 73 miles), 125 fathoms, one (S. Afr. Mus.
A8619. P.F. coll.).
Remarks. The Pieter Faure specimen came from almost the same locality
as the Valdivia specimen. The tip of the canal is broken, but otherwise the shell
is unworn. The thin, strongly crispate and outstanding plicae overshadow the
206 ANNALS OF THE SOUTH AFRICAN MUSEUM
spiral lirae, and thus the sculpture appears a little less distinctively cancellate
than in Thiele’s figure.
The smooth protoconch precludes the inclusion of this species in
Afritrophon.
Trophon mioplectos n. sp.
Fig. 44(a)
Protoconch 14 whorls, alt. 0:75, diam. 0-5 mm., smooth, junction with
Ist postnatal whorl clearly marked. Postnatal whorls 3; axial plicae on Ist
whorl 13, on 2nd 14, on 3rd 15, lamellate, sharp, from suture to suture and
extending across base to rostrum; profile of 2nd whorl obscurely biangulate,
of 3rd more distinctly biangulate. Tip of canal broken. No spiral sculpture.
4 Xx 2mm. Dirty white.
34° 24'S. 25° 42’ E., 256 fathoms, one (S. Afr. Mus. A8631. P.F. coll.).
Remarks. Differs from denseplicatus Turton 1932 and gemmulatus Turton
1932 (the latter appears to be synonymous with the former) in having fewer
axial plicae, presuming Turton’s count was correct. He gave ‘nearly 30° for
denseplicatus and ‘nearly 20° for gemmulatus. Both photographs are rather poor,
but even that of denseplicatus seems to indicate at most 20 plicae.
Trophon johannthiele: n. sp.
Fig. 44(6)
1925. Thiele. D. Tiefsee Exp., xvii, p. 170, pl. 30 (18), fig. 16 (Tvophon sp. juv.).
Protoconch 14 whorls, alt. 0:5, diam. 0:75 mm., smooth. Postnatal
whorls 3, profile biangulate, junction with protoconch abrupt. Axial plicae
11 on each whorl, strongly retractive from suture to shoulder and then vertical,
thin, with free edges, and squamosely raised at intersections with 2 feeble
spiral lirae. Plicae extend from suture to suture, and across base, with
squamulae on 2 additional lirae. Growth-lines obscure. Tip of rostrum
broken. 5 X 2°8 mm. Pale buff.
85119 ©. 20 12 120 meres: omnes (ihicle):
Off East London, 400-450 fathoms, one, exceedingly fragile (S. Afr. Mus.
A8844. P.F. coll.).
Remarks. ‘Thiele’s example with protoconch plus 2 whorls measured 3-6
(according to the magnification of his figure). ‘The thin free-edged plicae are
similar in this species and jucundus Th., but are more numerous and less strongly
squamate in the latter. Also the shape of the protoconch is different.
Trophon pistillum n. sp.
Fig. 44(/)
Protoconch 14 whorls, alt. 0-8, diam. 0:75 mm., smooth. Postnatal
whorls 3, profile angular, Axial plicae 11-12 on Ist whorl, 13-14 on and,
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 207
15-16 on 3rd, retractive (but less so than in johannthielez) from suture to shoulder,
then vertical, thin, with free edges, squamately raised at intersections with 2
feeble spiral lirae; plicae extending from suture to suture, and across base with
tubercles on 2 additional lirae. —The squamae on the plicae are not hollow in
front, but filled in with a tiny plug or tubercle, rather like the piston in a
cylinder (with incomplete rim). 3 or 4 growth-lines distinct between each
pair of plicae. Rostrum broken. Approximately 5 x 3 mm.
Operculum oval, nucleus apical.
Radula with c. 70 rows, central plate with 3 cusps, the median one larger
than the side cusps, postero-lateral corner of base acute.
Off Cape Natal (Durban), 440 fathoms, one, exceedingly fragile (S. Afr.
Mus. A8842. P.F. coll.).
Gen. AFRITROPHON Tomlin
1947. Tomlin. 7. Conch., xxii, p. 271.
Protoconch bicingulate. Postnatal whorls with one or more strong spiral
lirae; axial ribs present on Ist postnatal whorl; on following whorls either well
developed or obsolete. Sutures deep. Canal short. Operculum ovoid, nucleus
on outer margin near apex.
Radula, central plate with 3 rather large cusps, postero-lateral corners of
base acute, lateral plate slender.
Genotype: Trophon kowieensis Sow. 1901.
Remarks. Tomlin saw no specimen with an unworn protoconch, and did
not mention it in his generic diagnosis. Fortunately for his genus, however, he
happened to choose as genotype one of the species which has a peculiar proto-
conch.
Of the other four species included by Tomlin in his genus, insignis Sow.
and agulhasensis Th. have similar protoconchs, and are rightly included. But
jucundus ‘Th. and acutispira Sow. (if my identification of the latter is correct)
have a perfectly smooth protoconch and must be excluded. I refer them back
to Trophon.
Tomlin declined to comment on Turton’s (1932) four ‘species’. One
might, however, suggest that the figures of denseplicatus and gemmulatus indicate
merely different aspects of the same species; subglobosus seems to be a Turritella;
and ornatus is definitely the protoconch of a Turritid (cf. ‘Clathurella’ capensis).
An example of Thiele’s “Trophon sp. juv.’ has been found in the Preter
Faure material; as it has a smooth protoconch it also is retained in Trophon (see
johannthielei, p. 206).
Afritrophon kowieensis (Sow.)
Figs. 44(@), 45(@)
1901. Sowerby. Proc. Mal. Soc., iv, p. 213, pl. 22, fig. 16 (Trophon k.).
1925. Thiele. D. Tiefsee Exp., xvii, p. 169 (Trophon ? kowiensis typ. err.).
1947. Tomlin. loc. cit., p. 271.
208 ANNALS OF THE SOUTH AFRICAN MUSEUM
Protoconch 14 whorls, bicingulate. Postnatal whorls 4 (44); axial ribs
on Ist whorl 14-15, at first interrupted by the 2 spiral lirae, but later forming
nodules at the intersections, on 2nd whorl 18-20, on 3rd 26-28, on 4th c. 28-34,
from suture to suture, obsolete on base; on the 2nd whorl the nodules gradually
become vaulted squamae, hollow in front; on the last whorl they are closely
ageregated but vary in number; on grd and last whorls the axial ribs between
suture and upper peripheral lira also become vaulted squamae. On base 4-5
additional squamose lirae. 8 x 3:3 mm.
Operculum ovoid, nucleus on outer margin near apex.
Radula with c. 130 rows, as in generic diagnosis.
Kowie (= Port Alfred) (Sowerby, Bartsch, Turton).
Agulhas Bank 80 metres, St. Francis Bay, 80 metres, and Algoa Bay,
102 metres (Thiele).
Off Great Fish Point, 22 fathoms; Algoa Bay; off Cape Recife, 52 and
124 fathoms; off Knysna Heads, 46 fathoms; off Cape St. Blaize, 125 fathoms;
34° 27'S, 25 ° 42’ E., 256 fathoms (S. Afr. Mus. P.F. coll.).
Living: Algoa Bay, 60 fathoms; False Bay, 30 metres (U.C.T.).
Remarks. The original spelling was, quite correctly, kowie - ensis, and
should be retained.
At first sight somewhat resembling a very slender 7. scrobiculata (p. 212).
Recorded also by Thiele from Great Fish Bay, Angola, but the identity
of this specimen, should, with due respect to Thiele, be accepted with reserve.
Afritrophon agulhasensis (Thiele)
Fig. 45(¢)
1925. Thiele. D. Tiefsee Exp., xvii, p. 169, pl. 30 (18), fig. 12 (Trophon ? a.).
1947: Tomlin. loc: cit., p: 271-
Protoconch 14 whorls, bicingulate. Postnatal whorls 5 (Thiele gave total
number of whorls in his specimen 6); axial ribs on 1st whorl 15, at first inter-
rupted by the 2 spiral lirae, later forming nodules at the intersections; on 2nd
whorl 14-13, later whorls 13-12, nodules at intersections becoming stronger
and more outstanding, especially on the upper peripheral lira on 3rd—5th
whorls, some of them auriculate, hollowed in front; ribs extending from suture
to suture, obsolete on base; on base 4 less strongly nodose-squamose lirae and
an indistinct 5th on rostrum. 8 X 3°5 mm.
Operculum ovoid, nucleus on outer margin near apex.
Radula with c. 145 rows, as in generic diagnosis.
35° 10 §.,22° 26 E., 155 metres: ((hiele)s
Off Cape St. Blaize, 125 fathoms; 35°5’S. 25°55'’E., 67 fathoms;
34° 24'S, 25° 4a’ E., 256 fathoms (S. Afr. Mus. Pi. coll.)
Beach specimens: False Bay and Still Bay (S. Afr. Mus.).
Living: False Bay (U.C.T.).
————————
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 209
Remarks. Although similar to kowveensis, this species, as Thiele said, is
quite distinct: the area between suture and upper peripheral lira is quite
smooth except for the fine non-squamose axial plicae, and the axial plicae are
fewer; the nodules in unworn specimens are more outstanding and strongly
auriculate.
In Thiele’s specimen the grd (as here reckoned, = Thiele’s 5th) lira on
the base was not developed. .
Fic. 45.
(a) Afritrophon kowieensis (Sow.), apex and operculum. (6) A. insignis (Sow.). (¢) A. agulhasensis
(Thiele), with apex and protoconch further enlarged.
Afritrophon insignis (Sow.)
Fig. 45(4)
1900. Sowerby. Proc. Mal. Soc., iv, p. 2, pl. 1, fig. 1 (Trophon ? 7.).
1925. Thiele. D. Tiefsee Exp., xvii, p. 170, pl. 30 (18), fig. 14 (Trophon i.).
947°) lomlin. loc. cit., p. 271-
Protoconch 14 whorls, bicingulate. Postnatal whorls 4; 1st whorl with
axial ribs and 2 spiral lirae as in the two previous species; from end of Ist
whorl the lower lira is for the most part occluded by the suture of the following
whorl, though here and there a tiny auriculate tubercle projects; the upper
lira becomes very prominent with 8 or g sharp, auriculate, upturned tubercles;
on base the 2nd lira becomes visible again, with auriculate tubercles less
prominent than those on the upper lira; no other lirae on base; fine growth-
lines distinct. 6-5 X 3 mm.
Kowie (= Port Alfred) (Sowerby, Bartsch, Turton, S. Afr. Mus.).
Off Cove Rock (East London area), 22 fathoms, 3 dead but fresh (S. Afr.
Mus. A849. P. F. coll.).
Remarks. In worn specimens the projecting tubercles become merely
undulations on the peripheral keel; these undulations were not mentioned in
the original description, but can be seen in the cotypes in 8. Afr. Mus.
Thiele recorded a 3-7 mm. juvenile from Great Fish Bay, Angola; but I
feel that the identification needs confirmation. | |
210 ANNALS OF THE SOUTH AFRICAN MUSEUM
Gen. Typuis Montfort
1929. Thiele. Handbuch, i, p. 293, fig. 318 (radula).
Radula with wide central plate, 1-3 small cusps between the median and
side cusps, lateral plate unicuspid, uncinate.
Typhis arcuatus Hinds
Fig. 46(a)-(g)
1843. Hinds. Proc. Zool. Soc., Lond., p. 19.
1844. id. Moll. Voy. Sulphur, p. 10, pl. 4, figs. 1, 2.
? 1903. Von Martens. D. Tiefsee Exp., vii, p. 94, pl. 3, fig. 2 (transcurrens).
1925. Thiele. ibid., xvii, p. 170.
Protoconch 14 whorls, alt. 1-3, diam. 1 mm., smooth, glistening. Post-
natal whorls 5; 4 tubes and 4 varices on each whorl; tubes subcircular on early
whorls, becoming oval and carinate in front on later whorls, sometimes on
last whorl narrowly oval (complanate); varices curving forwards, carinate,
connected with the tubes, but with a shallow notch at base of tube defined by a
feeble angulation. 20 < 10-11 mm. (excl. tubes).
Operculum broadly oval, fitting the continuous peristome, nucleus
apical, growth-lines well marked.
Surface dull or chalky-white, except the glistening protoconch.
Fic. 46.
Typhis arcuatus Hinds. (a) body whorl. (6) anterior view of tube and varix. (c) the same of a
specimen from Cape Point (S. Afr. Mus. no. A4g45). (d), (e) diagrammatic apical views of
normal specimen, and Cape Point specimen. (f/f), (g) two views of protoconch. T. penta-
phasios n. sp. (h), (1) body whorl, and diagrammatic apical view.
————
“ te wt” Fo ene
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 211
Agulhas Bank, 40-45 fathoms (Hinds); 35° 16’ S. 22° 26’ E., 155 metres,
amd) 35° 19 5. 20° 12’ E., 126 metres (Thiele).
Off Cape Point, 123 fathoms; off Cape St. Blaize, 125 fathoms; off Cape
Recife, 124 fathoms, juveniles; off East London, 43 fathoms (S. Afr. Mus.
PP coll.).
Living: off Cape Point, 380-475 fathoms, one; off Cape St. Blaize,
55 fathoms, one (S. Afr. Mus. P.F. coll.).
Remarks. One specimen from off East London, although 5-whorled, is
smaller than the others (15 x 6 mm.), and appears proportionately narrower
because the tubes are curved upwards towards the apex; in typical specimens
they project obliquely upwards. In two examples from off Cape Point, 123
fathoms, the tubes on the last whorl are very broad (transversely), complanate,
and splayed outwards almost horizontally (fig. 46(c), (e)). The rather striking
appearance thus produced might seem to indicate a different species, but the
tubes become horizontally expanded only on the last whorl.
Five specimens have the varices carinate, with an obscure shoulder at the
top defining the shallow notch at base of tube. In the narrow East London
specimen the shoulder and notch are evanescent, and the last 3 varices become
progressively less carinate, in fact the last is broadly rounded. In another East
London example all the varices are broadly rounded, but those on the early
whorls have a faint carination on the hinder side.
The length of the tubes is no criterion as they are subject to wear and
corrosion.
Two juveniles with protoconch plus 14 whorls, 3-5 mm. long, and two
with protoconch plus 3 whorls, 7 mm. long., have been examined.
The animals of the two living Pieter Faure specimens were not preserved.
Apparently there is no character by which transcurrens, Zanzibar Channel,
5° 27'S. 39° 18’ E., 463-465 metres, can be separated except size: 6 whorls
(i.e. protoconch plus 5) 13 x 6 mm. (excl. tubes). Thiele, however, recorded
both species.
Sowerby compared. duplicatus (1870. Proc. Kool. Soc. Lond., p. 251, China)
with arcuatus, both species having curved varices.
Lyphis pentaphasios n. sp.
Fig. 46(/), (2)
Protoconch corroded. Postnatal whorls 4; 5 tubes and 5 varices on each
whorl; tubes oval-subcircular, with a broadly rounded rib below extending
to the suture; varices broadly rounded, midway between tubes and growth-
lines, profile evenly curved, not shouldered; peristome not quite continuous
where the varix from the tube impinges upon it. 11 (incl. corroded proto-
conch) X 5°5 mm. White.
Off Cape Point, 660 fathoms, two dead (S. Afr. Mus. A4g49. P.F. coll.).
212 ANNALS OF THE SOUTH AFRICAN MUSEUM
Tritonalia puncturata (Sow.)
Figs. 40(¢), 47(@)
1886. Sowerby. 7. Conch., v, p. 2 (Cominella p.).
1892. id. Mar. Sh. S. Afr., p. 11, pl. 1, fig. 9 (Cominella ? p.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 53, pl. 12, no. 393 (Cominella p.), and p. 53, pl. 12,
no. 394 (bipartita color. var.).
Protoconch 2 whorls, alt. and diam. 0-5 mm., smooth. Postnatal whorls 6.
Axial ribs on 1st whorl 15, on and and 3rd 15-16, on 4th 17-18, but becoming
fused one with another and obsolete on later whorls. Spiral lirae on tst whorl
3, on 2nd 4, on 3rd 5, on 4th and 5th 6—7, the lower 2 stronger than those above
and forming the peripheral shoulder, also usually a finer lira below the peri-
phery, on 6th whorl 5 (6) fine, 2 stronger peripheral, and 2 subperipheral lirae.
The sculpture on 1st and early part of 2nd whorl is cancellate, but thereafter
the spirals predominate, especially the two peripheral ones; the groove between
these two foveolate. On base 10-11 additional lirae, with punctate intervals.
Canal open. 20 (without protoconch) x 9 mm.
Operculum oval, nucleus near outer margin below middle.
Buff or brownish, with darker axial streaks, or darker above and below
the peripheral lirae, or dark above and pale below, or vice versa, lirae on body-
whorl often spotted.
Radula with c. 180 rows, central plate with median cusp arising near front
margin, 2 large submedian cusps on hind margin with 2-3 denticles between
each of these and the median cusp.
False Bay, Hermanus, Still Bay, Mossel Bay, Port St. Johns (S. Afr. Mus.).
Living: west coast of Gape Peninsula (S. Afr. Mus.); Jeffreys Bay (St.
Francis Bay), Knysna estuary, Mossel Bay, Breede River mouth (U.C.T.).
Remarks. The bicarinate and foveolate periphery is characteristic, as also
are the finer lirae above the strong peripheral lirae. Some variation may occur:
occasionally one of these finer lirae may be stronger than the others, almost as
strong as the peripheral lirae.
Juveniles (5 mm.), especially slender forms, before there is a clear
distinction between the fine upper lirae and the strong peripheral ones, are
liable to confusion with Trophon acutispira. The axial ribs and the additional
lirae on base are a little more numerous in puncturata.
Tritonalia scrobiculata (Dnkr.)
Fig. 47(4), (¢)
1846. Dunker in Philippi. Abbild. Beschr. Conch., ui, p. 118, pl. 3, fig. 4 (Fusus s.).
1892. Sowerby. Mar. Sh. S. Afr., p. 2 (Murex ? Ocinebra s.).
1892. id. ibid., p. 2, pl. 1, fig. 1 (Murex babingtoni).
1892. id. ibid., p, 2, pl. 1, fig. 2 (Murex crawford).
1932. Turton. Mar. Sh. Pt. Alfred, p. 75 (crawfordi) and p. 76 (scrobiculata and babingtoni).
Protoconch 14 (2) whorls, alt. 0:75, diam o-5 mm., smooth. Postnatal
whorls 4 (44). Axial ribs on 1st whorl 11-12, on and 12-14, on 3rd 15-16, on
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 213
last whorl c. 16-18 (but may occasionally be only 12); between suture and
upper peripheral lira growth-lines form more or less distinct pliculae cor-
responding with the squamous nodules on the lira, usually sinuous, and more
or less raised into vaulted squamulae; when strongly developed this series of
squamulae forms the first of the 3 spiral lirae characteristic of crawford: (fig.
47(¢)); sometimes there are 2 such lirae, or, on the 4th whorl 3, so that the
profile of 3rd whorl shows 4,.that of 4th whorl 5 spiral lirae (babingtoni). The
squamous nodules on the 2 strong peripheral lirae are very variable in strength.
The foveoles between the lirae are usually nearly square, but may be trans-
versely oblong when the number of ribs is reduced. On base 6 additional
lirae, the 4th usually the smallest.
Fic. 47.
(a) Tritonalia puncturata (Sow.), 5th whorl, and protoconch. (6) T. scrobiculata (Dnkr.), proto-
conch. (¢) 3rd and 4th whorls showing transition to 3-lirate crawfordi. (d) T. kienert (Rve.).
Sometimes the spiral lirae are scarcely thickened and only feebly nodulous,
consequently the axial ribs in the sulci are more distinct, and the sculpture
more obviously cancellate. Worn examples of babingtont appear distinctly
cancellate.
The outer lip may be thickened, with 4-6 denticles within at the margin,
but only when fully adult. Canal open. 15 x 7°5 (8:5) mm.; Turton: 22 mm.
e typ. err.
Operculum oval, nucleus near outer margin below middle.
Buff or brownish. :
Radula with c. 150 rows, similar to that of puncturata.
Table Bay and west coast of Gape Peninsula, False Bay to Port Alfred, and
Tongaat (Natal) (S. Afr. Mus.).
Off Knysna, 46 fathoms; off Keiskamma River, 33 fathoms; off East
London, 32 fathoms (S. Afr. Mus. P.F. coll.).
Living: Langebaan (Saldanha Bay), and False Bay (U.C.T.).
Soa ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. A common and distinct but variable species, which, it is not
surprising, has been described from beach-worn specimens under three separate
names.
Tritonalia purpuroides (Dnkr. MS.) (Rve.)
Dunker. MS. (as var. of scrobiculatus).
1845. Reeve. Conch. Icon., Murex no. 158, pl. 32.
1848. Krauss. Siidafrik. Moll., p. 112, pl. 6, fig. 14 (Murex dunkert).
1903. Von Martens. D. Tiefsee Exp., vii, p. 25 (Murex p.).
1925. Thiele. ibid., xvii, p. 168 (Murex p.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 76.
Protoconch 14 whorls, alt. and diam. 0-5 mm., smooth. Postnatal whorls
4-5. Axial ribs on 1st whorl 11, on 2nd 11-12, on 3rd (13) 12-11, on 4th
10-9 (8), on 5th 9-8. Spiral lirae 2 strong ones forming the peripheral shoulder;
on base 6—7 additional lirae, often between the lower peripheral and the 1st
basal lirae a thin lira, and often a similar one between 1st and 2nd basal lirae.
Space between suture and upper peripheral lira smooth except for the fine
growth-lines and axial plicae, the latter never squamulose; sometimes indica-
tions of 2 (very occasionally 3) fine spiral lirae between, but not crossing, the
axial plicae. Intersections of axial ribs and peripheral lirae nodulous, or
raised into pseudo-squamae (not or scarcely vaulted or hollowed in front).
Basal lirae broader than the intervals, often very broad so that the intervals
are mere striae. Outer lip in adult thickened, with 6—7 denticles or plicae
within. Canal open. 15 X 7 mm.
Beach examples: white, often a faint brown band around middle of
whorl, best seen within the aperture.
Cape, Port Elizabeth (Krauss, Bartsch, Sowerby); Port Alfred (Turton:
only one specimen). Agulhas Bank, 35° 29'S. 21° 2’ E., 102 metres, and
34° 51'S. 19° 37’ E., 80 metres (von Martens).
Dassen Island, west coast of Cape Peninsula, and Kalk Bay (False Bay)
(S. Afr. Mus.).
Remarks. Has not been taken alive. Apparently does not occur farther
east than Port Elizabeth, though Turton claimed to have found one specimen
at Port Alfred.
With fewer axial ribs than scrobiculata, especially on the last whorl. Most
specimens, especially the extreme forms of the two species, are easily separable;
but occasionally a specimen occurs which appears transitional and difficult to
assign to one or the other. The absence, however, of squamulae between the
suture and the shoulder lira seems to be a constant feature of purpuroides.
Tritonalia kienert (Rve.)
Fig. 47(¢)
1845. Reeve. Conch. Icon., iii, Murex sp. 172 (Murex k.).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 60 (Tritonalia k.).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 215
1925. Thiele. D. Tiefsee Exp., xvii, p. 168, pl. 30 (18), fig. 9 (Murex k.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 76.
Protoconch 2 (24) whorls, alt. 1, diam. o-8 mm., smooth. Postnatal
whorls 4; axial ribs on Ist whorl 10-11, on 2nd 11, on 3rd and 4th 10-11 (3rd
whorl in two slender specimens with only 9g ribs), from suture to suture and
extending across base, including the groove, to rostrum; spiral lirae one peri-
pheral on first half, 3 on later half of 1st whorl, 3-4 on and whorl, 4 on 3rd
and 4th whorls, on base 3 additional lirae followed by a deep groove, then 2
nodose lirae on rostrum; this groove forming a notch in outer lip is already
developed on the 2nd whorl in the smallest specimen examined 4:5 mm. long.
Canal narrow but not closed. 18 mm. (Turton). 12-13 mm. with worn apices
(S. Afr. Mus.). Plump and siender examples (with 3 whorls): 8 x 4, 9 X 4,
9 X 4°5 mm.
Natal and Algoa Bay (Sowerby); Port Alfred (Bartsch, Turton); St.
Francis Bay, 80 metres (Thiele); Still Bay and False Bay (S. Afr. Mus.).
Tritonalia sperata (Cossm.)
Fig. 48
1904. Smith. 7. Malac., xi, p. 30, pl. 2, fig. 11 (Fusus cingulatus, non Sowerby 1832).
1921. Cossmann. Rev. Crit. Paléozool., xxv, p. 181 (Fusus s.).
1931. Tomlin. Ann. Natal. Mus., vi, p. 434 (Fusinus s.).
1933. Turton. Mar. Sh. Pt. Alfred, p. 50 (Fusus speratus Crossman [sic] typ. err.).
1957. Barnard. 7. Conch., xxiv, p. 180, fig. (radula) (Latiaxis fritschi, non von Martens).
1959. id. ibid., xxiv. p. 327.
Protoconch 2 whorls, alt. 1-1-25, diam. 1:3-1:5 mm., smooth, junction
with Ist postnatal whorl marked by several fine pliculae. Postnatal whorls 5,
profile convex, in later whorls bluntly shouldered; axial ribs on 1st whorl
15-14, on 2nd 14-13, on 3rd 13-12, on 4th 11-10, on 5th 9-8, from suture to
suture until 3rd whorl, thereafter petering out above and below the periphery
where they form blunt rounded knobs, especially prominent on the body-
whorl; crossed by spiral lirae 3 on Ist whorl (uppermost feeble at start),
increasing to 4-6 on 2nd whorl, and very numerous on later whorls, c. 18-20
on 5th whorl; on 3rd whorl peripheral lira usually thicker than the others and
- rather prominent; on base c. 25 additional lirae (some being very fine inter-
mediaries), on upper part of base one lira (usually the 5th) stronger than the
others, forming a costa, rostrum costate; all the lirae minutely squamulose.
Columella curved, its edge free and anteriorly produced over, but not com-
pletely closing the canal. 24 (apex missing, only 3rd—5th whorls present) x
Bim. Smith: 28 xX 14 mm. .
Operculum oval, nucleus on outer margin below middle.
Grey with a rosaceous or violaceous tinge (U.C.T. specimen); fawn or
buff (P.F. specimens, several years in alcohol). Beach specimens pinky-
orange or salmon, the protoconch more deeply coloured.
216
ANNALS OF THE SOUTH AFRICAN MUSEUM
Radula with at least 340 rows, central plate with median cusp arising
from front margin, not extending as far as side cusps, which have one denticle
on inner margin and 2-3 externally, postero-lateral corner of base acute,
lateral plate moderately stout.
Dead: Port Alfred (Smith, Bartsch, Turton, S. Afr. Mus.).
Living: off Gape Morgan, 36 fathoms; off East London, 34-47 fathoms;
Mossel Bay, 16 fathoms (S. Afr. Mus. P.F. coll.). Mossel Bay (U.C.T.).
Fic. 48.
Tritonalia sperata (Cossm.).
Remarks. The costa on base (Smith: infraperipheral) is scarcely visible on
3rd whorl, feeble on 4th, and only becomes well developed on the 5th, and even
then is variable in strength.
1919.
T7715
1816.
1822.
1848.
1859.
1919.
Gen. Tuais Bolten
Cooke. Proc. Mal. Soc., xiii, p. 91, figs. (radulae).
Thais cingulata (Linn.)
Fig. 49(/)
Linne. Mantissa, i, p. 549 (Buccinum c.).
Lamarck. Tabl. Encycl. Meth., pl. 422, figs. 4a, 6. and Liste, p. 5 (Triton trochlea).
id. Anim. sans Vert., vii, p. 248 (trochlea and clavus).
Krauss. Stidafrik. Moll., p. 118.
Chenu. Man. Conchyl., i, fig. 805.
Cooke. loc. cit., p. 94, fig. 12 (radula).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 217
Protoconch 14 whorls, alt. 1, diam. 1-1-25 mm., smooth. Postnatal
whorls 5, 1st demarcated from protoconch; axial ribs on 1st whorl 19-20,
closer together at start than on later part of whorl, on and and grd whorls
18-20, thereafter becoming irregular and intermixed with the finer growth-
lines; spiral keels 2 on first part of 1st whorl, becoming 3 on later part, 3 on
2nd whorl, the upper one forming a shoulder girdle, the middle one peripheral,
the lower one smaller and forming a suprasutural lira often concealed by the
succeeding whorl; 3rd whorl with typically 3 keels, 4th and 5th with typically
3 or 4 (4 keels may develop sometimes on later part of 3rd whorl) ; additional
lirae may develop on 4th and 5th whorls (sometimes on later part of 3rd)
between suture and the shoulder keel, between each pair of keels, and also on
base, or in the extreme non-cingulate form over the whole whorl and all whorls
except the 1st. Outer lip internally plicate or grooved (or both), corresponding
with the external girdles and the presence or absence of lirae. 41 X 23 mm.
(unicingulate); 35 X 21 mm. (tricingulate); 20 x 13 and 20 X IO mm.
(typical). Smallest examples seen (protoconch plus 2 whorls) 4-5-5 mm. long.
Operculum oval-oblong, nucleus at outer margin, without distinct trans-
verse oval lines.
Fic. 49.
(2), (b) Thais squamosa (Linn.), with subscalariform aberration. (c) T. capensis (Petit). (d) T.
wahlbergi (Krss.). (e) 7. dubia (Krss.). (f) T. cingulata (Linn.). (g) T. castanea (Kiister).
218 ANNALS OF THE SOUTH AFRICAN MUSEUM
Grey, buff, or dirty white, operculum horny.
Radula with c. 165 rows (shell 10 mm.), ¢. 310 rows (shell 30 mm.),
central plate with median cusp longer than side cusps, latter with one denticle
on inner margin and 1-2 externally, postero-lateral corner of base acute,
lateral plate rather slender.
Table Bay and Dassen Island; False Bay (S. Afr. Mus.).
Living: Port Nolloth (U.C.T.).
Remarks. Krauss gave Natal as well as the Cape as the habitat, but except
for Sowerby’s (1892) solitary record from Port Elizabeth, there are no records
and no examples in 8. Afr. Museum from any locality east of False Bay.
Dr. Muir did not find it at Still Bay. The Port Elizabeth record is probably
another instance of ‘domicile of the collector, not of the mollusc’.
The axial ribs are entirely subordinate to the spiral keels on the 2nd and
following whorls, but strongly enough developed on 2nd and 3rd whorls to
form a clathrate sculpture. On the early whorls the keels are slightly nodular
at the intersections, and on later whorls successive major growth-lines (or
temporary stoppage of growth) are indicated by slightly irregular junctions.
The variable development of the girdles on the later whorls is even more
striking in this species than in Burnupena cincta (p. 160). Krauss stated that the
girdles varied from 1-4, and sometimes there were none at all. Possibly this
statement has acted as a warning to later authors not to give varietal or specific
names to the several forms; the extreme forms—5-—cingulate (not mentioned by
Krauss) and non-cingulate—are certainly very different in appearance.
When well developed (and unworn) the girdles are often very strong, with
their upper and lower margins curling over; they may vary in width; and they
may also be nearly flat, with little relief above the general surface of the shell,
thus grading into the non-cingulate form. When the girdles are suppressed,
the lirae are better developed. The non-cingulate form is wholly lirate, the
lirae being equally regular in width throughout, or the girdles may be just
indicated by slightly wider lirae. In some examples the lirae supersede or
obscure the bicingulate and clathrate sculpture on the 2nd whorl which is
characteristic of the typical form.
The three hundred (308) available specimens (most of them from a
private collection, and probably from Table Bay or the west coast of the Cape
Peninsula) seem to show that the tricingulate form is the commonest (185
specimens), the quadricingulate the next commonest (91); there are 12 bicingu-
late, 8 unicingulate, 6 non-cingulate, and 4 (all large) quinquecingulate. ‘Two
others are tricingulate, but the later part of the 4th whorl (? due to injury)
suddenly loses one girdle, in one case the Ist, in the other the 3rd girdle. ‘The
12 bicingulates include eleven with 1st and 2nd girdles, but one with 2nd and
3rd girdles; the 8 unicingulates include 2 with the 2nd girdle and 6 with the
ist girdle. There are no unicingulate examples in which the only remaining
girdle is the 3rd or 4th.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 219
The non-cingulate form has some similarity with wahlbergi, but is distin-
guished by smaller protoconch, the bicingulate 2nd whorl, and the absence
of axial ribs on later whorls.
It also resembles even more closely some froms of dubia (e.g. Oudekraal,
p- 223), but the protoconch and 1st whorl, and the whole spire, are broader.
Krauss (loc. cit., p. 118) recorded succincta from the Cape, but probably
Baron von Ludwig’s collectign contained shells from various localities and
possibly not always with locality labels. Lamarck’s figures (1816. abl. Encycl.
Meth., pl. 398, figs. 1a, 1b) show 8 girdles on the body whorl. Sowerby (1802,
p- 13) said the species lives at Madagascar and other eartern localities.
+ Thais praecingulata (Tomlin MS.) Haughton
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 48, pl. 5, figs. 6-10.
Fossil, Pleistocene: Alexander Bay, Namaqualand; and Rietvlei,
Bredasdorp.
A large species: 95 (apex missing) X 65 mm.
Thais squamosa (Lam.)
Fig. 49(a), (2)
1816. Lamarck. Tabl. Encycl. Meth., pl. 398, figs. 2 a, b., and Liste, p. 2 (Purpura s.).
1832. Blainville. Nouv. Arch. Mus. Paris, i, p. 251, pl. 12, fig. 6 (Purpura clathrata).
1903. Smith. Proc. Mal. Soc., v, p. 376 (says clathrata is juv.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 78, also p. 78, pl. 18, no. 565 (sculpturata, the right-
hand figure, ? the left-hand figure).
Protoconch 2-24 whorls, alt. and diam. 1 mm. (sometimes alt. 1:3 mm.),
smooth. Postnatal whorls 6, 1st sharply demarcated from protoconch. Axial
ribs on 1st whorl 18-19, on 2nd 19-20, increasing to 22-25 or more on 5th
whorl, where they become confused with the growth-lines, from suture to suture
and, extending across base; spiral lirae on 1st whorl 2 (3 towards end of whorl,
or when not covered by 2nd whorl), on and 3-4, on 3rd 4-5, on 4th and 5th 5,
with intermediaries, on base 7 additional lirae with intermediaries, the 3rd
(sometimes 2nd) strong and often forming a costa on the body-whorl; inter-
sections of ribs and lirae nodular, on later whorls the nodules transversely
oblong, often subsquamose and hollowed in front. Rostrum squamose; aperture
(at certain stages of growth) plicate internally. Periostracum with short
spiniform processes. 61 (protoconch and 2 whorls missing) x 40 mm.;
44. X 27 mm.; 39 X 24 mm. Protoconch plus rst whorl 2-5 mm. long.
Operculum oval, nucleus on outer margin below middle.
Brown, uniform or with darker axial streaks, sometimes whitish with
brown spiral bands, or bright brown markings and axial streaks.
Radula with c. 190 rows (shell 20 mm.), ¢. 285 rows (shell 44 mm.) central
plate with median cusp not much longer than side cusps, latter with a denticle
220 ANNALS OF THE SOUTH AFRICAN MUSEUM
on inner and outer side, margin external to side cusp with 2-3 denticles,
lateral plate moderately stout.
Cape and Natal (Krauss); Port Elizabeth and Port Alfred (Sowerby,
Bartsch, Turton) ; ‘Table Bay and Dassen Island (S. Afr. Mus.).
Living: Lambert’s Bay, False Bay, and 34° 7’ S. 25° 46’ E., 41 fathoms
(U-GIT.). ‘Stall Bay *(S: Ate) Wias.).
Remarks. Plump and slender forms occur: 26 x 16 mm., and 27 X 14mm.;
52 X 35 mm., and 52 x 28 mm.
One specimen, 45 X 30 mm., has unusually strong spiral lirae and axial
ribs; this is particularly noticeable on the 4th and the beginning of the 5th
(body) whorl, producing a very strong clathrate sculpture with deep inter-
vening pits.
The protoconch sometimes sits very prominently on the first whorl.
Thais capensis (Petit)
Figs. 40( f), 49(¢)
1848. Krauss. Stidafrik. Moll., p. 117 (luteostoma, non Desh.).
1852. Petit. 7. de Conchyl., iii, p. 162, pl. 7, fig. 6 (Purpura c.).
1892. Sowerby. Mar. Sh. S. Afr., p. 14 (? var. of luteostoma).
1903. Smith. Proc. Mal. Soc., v, p. 376, pl. 15, fig. 21 (Purpura pura).
1904. id., 7. Malac., xi, p. 32, pl. 2, fig. 15 (Purpura texturata).
1919. Cooke. loc. cit., p. 93 (radula).
1923. Tomlin. 7. Conch., xvii, p. 47 (pura and texturata = capensis juv.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 76, pl. 18, no. 554, p. 77 (var. luteostoma Desh.), p. 77,
pl. 18, no. 556 (albolineata), and p. 77 (texturata).
1934. Nardini. Palaeontogr. Ital., 34 (1933), p. 205, pl. 15 (2), figs. 15a—-d (Purpura succincta
Mart. var. natalensis).
Protoconch 14-2 whorls, alt. 1°75, diam. 2 mm., smooth. Postnatal
whorls 5, 1st not clearly demarcated from protoconch. Spiral lirae on ist
whorl 7, the 5th stronger than the others, on 2nd whorl 10, the 6th and roth
stronger, 6th in middle of whorl, roth more or less concealed by succeeding
whorl (but visible on body-whorl); on 3rd and following whorls lirae increase
in number by interpolation; 2 additional strong lirae on base; from 3rd whorl
the 2 peripheral lirae become gradually nodose, 9-11 nodules, on last whorl in
large specimens 8-9 large blunt nodules, those of the upper series alternating
with those of the lower series; growth-lines in interstices between lirae well
marked (in fresh specimens) forming a punctate-striate sculpture. 60 (proto-
conch missing) * 34 mm., 55 (protoconch and 1st whorl missing) X 31 mm.,
51 (protoconch and first 3 whorls missing) x 34 mm. (knobs excluded in
measurements of width); 41 (incl. protoconch) x 23 mm. Protoconch plus
1st whorl 4°5—5 mm. long.
Operculum oval-oblong, nucleus on outer margin below middle, without
distinct transverse oval lines. |
Buff, brown, grey, the peripheral lira (lirae) with brown spots correspond-
ing with the knobs, aperture yellowish with brown bands corresponding with
~<.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 22
the spaces between the external lirae (or castaneous brown with 4 yellow
stripes); worn shells brown or yellowish, with brown spots of the lirae; proto-
conch whitish, 1st and 2nd whorls sometimes with brown axial streaks; oper-
culum horny-brown. One specimen (S. Afr. Mus.) uniform pale buff, almost
white, operculum horny-brown.
Radula with c. 95 rows (shell consisting of protoconch plus 1st whorl), at
least 160 rows (shell 36 mm.), central plate with median cusp much stronger
than side cusps, latter with or without denticle on inner margin, externally no
denticles or wrinkles, lateral plate rather stout.
Natal (Krauss, Smith, S. Afr. Mus.); Port Alfred (Smith, Bartsch, ‘Turton,
S. Afr. Mus.); Still Bay (S. Afr. Mus.). Cape Agulhas, collected by Verreaux
(Petit).
Living: Scottburgh, Natal (Cooke). Off East London, 195 fathoms
(S. Afr. Mus. P.F. coll.). Mossel Bay and East London (U.C.T.).
Remarks. In 1886 Sowerby did not agree with Tryon in regarding capensis
as a variety of the Japanese /uteostoma; but in 1892, having seen ‘an undoubted
luteostoma from Port Elizabeth, he was doubtful and remarked that the typical
forms were very unlike. As the original provenance of the Port Elizabeth
luteostoma is very doubtful, and cannot be checked, the Japanese species should
be excluded from the South African fauna-list. According to Cooke (1919)
there are slight differences in the radulae of the two species. My specimen of
capensis confirms Cooke’s description.
The protoconch is a little larger than that of Burnupena cincta and the lirae
on ist portion (or first portion of Ist postnatal whorl) are not so regular in
size and spacing, the 5th and 7th lirae being slightly thicker; very fine growth-
lines foreshadow the sculpture on the later whorls.
Turton’s albolineata seems to be merely a very worn specimen.
Nardini had one specimen, 25 < 16 mm., from Umkomaas. Fortunately
he gave an enlarged figure (photographic) of the sculpture, which puts the
identity with capensis beyond doubt.
Thats dubia (Krauss)
Figs. 40(g), 49(¢)
1836. Kiener. Cog. viv., pl. 40, fig. 94 a (Purpura lagenaria var.).
1848. Krauss. Stidafrik. Moll., p. 117.
1889. Sowerby. 7. Conch., vi, p. 148 (scobina, non Q. & G.).
1892. id. Mar. Sh. S. Afr., p. 14. (cataracta, non Chemn.).
1910. Schwarz. Tr. Geol. Soc. S. Afr., xii, p. 114 (Purpura scobina).
1919. Cooke. loc. cit., p. 93 (radula) (cataracta, non Chemn.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 77 (cataracta and cataracta dubia).
1932. id. ibid., p. 77, pl. 18, no. 561 (pyramidalis = juv.), and p. 78, pl. 18, no. 562
(rufanensis = juv.).
1932. Haughton. Tr. Geol. Soc. S. Afr., xxxiv (1931), p. 47.
Protoconch 14 whorls, alt. 1, diam. 0-75 mm., smooth. Postnatal whorls
5, Ist marked off from protoconch by a fine plica, or indicated only by the
222 ANNALS OF THE SOUTH AFRICAN MUSEUM
start of the spiral lirae and the growth-lines. Axial ribs on 1st whorl 15-16,
feeble at first and not always distinct from the growth-lines, on 2nd whorl
13-14, on 3rd 11-12; the ribs entirely subordinate to the spiral lirae but
forming nodules or vaulted scales or undulations on the lirae at the inter-
sections, usually not continued across base, on later whorls not distinct from
the growth-lines; spiral lirae 2 on ist, gnd and 3rd whorls, rather far apart,
becoming obscure, but usually traceable in good specimens on 4th and 5th
whorls, 4—5 additional lirae on base; from 3rd whorl onwards feeble inter-
mediaries may develop: 1 or 2 between suture and rst lira, one between the 2
peripheral lirae, and one between each pair on the base. 43 (protoconch and
2 whorls missing) x 28 mm., 41 (protoconch and 3 whorls missing) x 28 mm.,
24 X 14°5 mm., 22 < 16-5 mm., 20 X 15 and 20 X 11 mm. Protoconeamias
Ist whorl 1-75—2 mm. long.
Operculum oval-oblong, nucleus on outer margin, transverse oval lines
(thickenings on internal surface) distinct.
Typically pale with dark brown or blackish axial flames from suture down-
wards, making spots on the lirae, in larger specimens becoming zigzag markings
across nearly whole of body-whorl but leaving a more or less clear band around
middle. Considerable variation in colour and pattern occurs. There may be
dark disconnected irregular blotches, or spiral streaks only; or pale grey or
buff with faint yellowish spiral bands; or orange-brown with chestnut spiral
bands. Aperture castaneous with a pale band in middle, more distinct in
adults than in juveniles; even in the pale almost unicolorous examples where
the aperture is only suffused, the pale band is distinguishable though faint;
the columella also is more or less suffused. Examples from Langebaan
(Saldanha) have a very dark, almost purplish, aperture and columella.
Radula with c. 95 rows (shell 13 mm.), c. 270 rows (shell 29 mm.), central
plate with median cusp longer than side cusps, latter with a denticle on inner
and usually also on outer margin, externally one denticle or none, postero-
lateral corner of base pointed but not prominent, lateral plate rather stout.
Fossil, Quaternary: Namaqualand (Haughton); Sedgefield near Knysna
(Martin); Port Elizabeth (Schwarz).
Cape (Krauss) ; Port Elizabeth (Sowerby) ; Port Alfred (Bartsch, Turton) ;
Table Bay, False Bay, Still Bay (S. Afr. Mus.).
_ Living: Lambert’s Bay, Langebaan (Saldanha Bay), west coast of Cape
Peninsula, False Bay, Knysna (U.C.T.).
Remarks. Juveniles are slender with sharply pointed spire (pyramidalts) ;
this slenderness may be maintained in later whorls, but is usually succeeded
by plumpness.
Young specimens with well-marked nodules or vaulted scales on the lirae
are different in appearance from smoother adults. An 18 mm. example from
sheltered water in Knysna estuary (U.C.T.) has all the growth-lines very well
marked with free edges, and the temporary outer lips form squamate pro-
jections on each lira.
14
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 223
Sculpture and coloration are often obscured by corrosion and algal
growth.
Specimens collected at Oudekraal (west coast of Gape Peninsula) (U.C.T.,
also K.H.B.) with 4 whorls, 16-17 mm. long, are uniform pale buff or greyish-
white, with pale violaceous columella and aperture, the pale internal band
faintly visible. Their chief peculiarity, however, is the nearly equal size of the
main and intermediary lirae, especially on 4th whorl; the gnd and 3rd whorls
might be described almost as lirate and the body-whorl of one of them is
definitely quadrilirate, with 9 lirae on base. One of them fortunately retains
the prominent protoconch and narrow bilirate 1st whorl, and thus (together
with the pale band in the aperture) indicate their specific identity. The
operculum has distinct transverse oval lines.
A specimen in 8. Afr. Mus. (Reg. no. 6620. ‘South Africa’) was identified
by J. H. Ponsonby as the New Zealand Purpura scobina Q. & G., a species
recorded by Sowerby from Port Elizabeth. This specimen, however, is merely
a young dubia. Sowerby’s identification was also probably wrong, and Haustrum
scobinum should be deleted from the South African fauna-list.
Turton’s pyramidalis is obviously a young dubia. Turton evidently did not
realize the slenderness of this species when young; he gives the number of post-
nuclear whorls as 6, which seems to be either a misprint or a miscount: a
specimen 15 mm. long should have 4 postnatal whorls. His rufanensis is a squat
form of dubia (with apex worn).
Thais wahlbergi (Krss.)
Figs. 40(h), 49(d)
1848. Krauss. Stidafrik. Moll., p. 118, pl. 6, fig. 15 (Purpura w.).
1891. Smith. Proc. Zool. Soc. Lond., p. 436, pl. 4, fig. 2 (Coralliophila w.).
1892. Sowerby. Mar. Sh. S. Afr., p. 4 (Euthria w.).
1938. Bright. Trans. Roy. Soc. S. Afr., xxvi, p. 62 (Thais w.).
1947. Stephenson. Ann. Natal Mus., xi, pp. 273, 274 (Cominella w.).
1956. Orr. Proc. Ac. Nat. Sci. Philad., cviii, p. 250 (‘Purpura’ w.).
Protoconch 2 whorls, alt. and diam. 1-3—1-5 mm., smooth, prominent when
not worn. Postnatal whorls 54—6, profile convex, not or very feebly shouldered,
1st whorl sharply demarcated from protoconch. Axial ribs often obscure, on
Ist whorl 14, on 2nd 12, on grd 12-11, on 4th 11-10, thereafter irregular and
feeble, from suture to suture, obsolete on base even in juveniles; spiral lirae on
Ist and 2nd whorls 3 (on 2nd whorl one or two feebler ones below suture),
increasing by interpolation on later whorls, broader and narrower lirae more
or less regularly alternating, or 2 fine lirae between a pair of broader ones, the
intervals becoming very narrow, merely striae; when fresh and unworn the
close-set lines of growth make the lirae crispate (Smith) or finely scabrous, and
the striae punctate. Sculpture cancellate (transversely oblong hollows) on first
2 whorls, on later whorls spirally lirate/striate, with only feeble and irregularly
spaced axial ribs. Rostrum rimate. Canal when fully developed nearly closed,
224 ANNALS OF THE SOUTH AFRICAN MUSEUM
Outer lip sometimes thickened with a varix, internally plicate. 47 * 21 mm.
Protoconch plus 2 whorls 5:5 mm.
Operculum oval, nucleus on outer margin near apex, oblique (not trans-
verse) oval thickenings across centre of inner surface, not visible externally.
Grey or dirty whitish, operculum pale horny.
Radula very long, at least 360 rows, central plate with median cusp
arising on front margin, side cusp with a denticle on inner margin and one or
two denticles externally, lateral plate rather slender.
Animal cream-coloured, eyes very small, black.
Living: off Sea Point (Cape Town) (Smith; also S. Afr. Mus.); Oudekraal
(west coast of Cape Peninsula), Langebaan (Saldanha Bay), and Port Nolloth
(U.C.T.). Saldanha Bay (Orr).
Remarks. The fine crispate lirae, with intervening punctate striae, make a
characteristic sculpture.
Plump and slender forms occur: 43 X 23, 46 X 22, 46 X 20 mm.
Apparently a west coast species. Krauss’s locality ‘Natal’ has not been
confirmed, and there are no records from any intermediate localities.
Sowerby put this species into Huthria in spite of Krauss saying the oper-
culum was Purpuroid. According to the radula it can be included in Thais.
Orr has suggested in or near Urosalpinx.
Thais castanea (Kiister)
Figs. 40(2), 49(g)
1886. Kiister. Conch. Cab., p. 170, pl. 28, figs. 8, 9 (Purpura c.).
1886. Sowerby. 7. Conch., v, p. 3 (Cominella unifasciata).
1892. id. Mar. Sh. S. Afr., p. 11, pl. 1, fig. 11 (C. unifasciata).
1897. id. Append. Mar. Sh. S. Afr., p. 4 (C. unifasciata var. concolor.).
1938. Eyre and others. Ann. Natal Mus., ix, p. 110 (Cominella c.).
1938. Peile. Proc. Mal. Soc., xxiii, p. 99, fig. 35 (radula). |
Protoconch 14 whorls, alt. and diam. 0-75 mm., smooth. Postnatal whorls
5, start of 1st whorl distinct. Axial ribs on 1st whorl 10-11, but not very
distinct, on 2nd and following whorls 10 (sometimes only 9), forming distinct
blunt knobs at intersections with spiral lirae, more distinct on the upper than
on the lower lira, often obscure on last part of 5th whorl, in fresh specimens the
knobs are squamate, hollow in front; ribs not crossing base; spiral lirae on Ist
and following whorls 2, the upper some little distance from the suture, the
lower just above and often partially obscured by the succeeding whorl, some-
times a fine intermediary between the two; on base 4 additional lirae, often
more distinct in juveniles than adults, sometimes with fine intermediaries;
growth-lines producing a very finely striate appearance. Outer lip plicate
internally. 15 X 7:5 mm. Protoconch plus 2 whorls 2:5 mm. long. |
Operculum oblong-oyal, nucleus on outer margin, no distinct transverse
oval lines, |
CONTRIBUTIONS TO KNOWLEDGE OF 58.A. MARINE MOLLUSCA 225
Castaneous-brown, usually with the protoconch and ist whorl, or 1st and
2nd whorls, white; or brown with a white band in middle of body-whorl; or
white with a brown band below the suture; or uniform white.
Radula with c. 130 rows, central plate with median cusp larger than side
cusp, latter with denticle on inner margin, externally 2-3 denticles, lateral
plate stout. Peile said the radula was shorter than those of many species of the
family (Thaididae), only 88 rows. His examples were probably young.
Cape (Kuster), Natal (Sowerby), Port Alfred (Smith, Bartsch, Turton) ;
Pondoland and Still Bay (S. Afr. Mus.).
Off Tugela River (Natal), 65-80 fathoms, one dead but fiesh; off East
London, 45 fathoms, dead but fresh; off Sandy Point (north of Kei River),
51 fathoms (S. Afr. Mus. P.F. coll.).
Living: East London, and Still Bay (U.C.T.).
Thais distinguenda (Dnkr.)
1852. Dunker. Reise Novara. Moll., figs. 1-3.
1880. Von Martens. Mauritius G Seychellen, p. 236 (as var. of hippocastanum).
1919. Cooke. loc. cit., p. 95, fig. 4 (radula) (¢ntermedia, “usually regarded as var. of hippo-
castanea’ [sic]).
1952. Satyamurti. Bull. Madras Govt. Mus., n.s. I, 2, pt. 6, p. 167, pl. 16, fig. 5 (intermedia).
1952. Braga. Anais Est. zool. Ultramar., vii, 3, pl. 3, fig. 4 (as Ricinula tuberculata Blainv.).
Radula with c. 250 rows (shell 28 mm.) to 290 rows (shell 38 mm.),
central plate with median cusp longer than side cusps, latter with denticle on
inner margin, 3 denticles or wrinkles externally, lateral plate moderately stout.
Living: Umpangazi (Zululand) (U.C.T.). Delagoa Bay and Mozambique
Island (K.H.B. coll., and U.W.). Ponta do Ouro (Mozambique) (Braga).
Distribution. Nicobars, Mauritius, Indo-Pacific.
Remarks. The U.C.T. specimens came to me labelled intermedia (Kiener).
Mr. Salisbury has identified a specimen as distinguenda (var. savignyi Desh.) ;
and the specimens are here recorded under this name. Braga’s illustration
seems to agree with these specimens.
Von Martens regarded distinguenda as synonymous with hippocastanum
(Linn.) var. intermedia (Kien.). He mentioned the variation in colour of the
columella: dark with a white stripe (intermedia), brownish with paler patches
(hippocastanum s.s.) whitish (savigny). Some of the present specimens resemble
hippocastanum, but most of them correspond better with savignyv.
Thais bitubercularis (Lam.)
1822. Lamarck. Anim. sans. Vert., vii, p. 237 (Purpura b.).
1836. Kiener. Cog. viv., p. 49, pl. 11, fig. 32.
1859. Chenu. Man. Conchyl., i, fig. 803.
1919. Cooke. loc. cit., p. 92 (radula).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 169, pl. 7, figs. 7 a, 6.
Radula with c. 195 rows, central plate with median cusp much stronger
and longer than side cusps, a separate small denticle between median and side
226 ANNALS OF THE SOUTH AFRICAN MUSEUM
cusps, externally 2 minute denticles or wrinkles, lateral plate rather stout.
Living: Delagoa Bay (U.W.).
Distribution. Karachi (Cooke). East Indies.
Thais gemmulata (Lam.)
1764. Linne. Mus. Ulricae, p. 636 (mancinella part).
1816. Lamarck. Tabl. Encycl. Meth., p. 397, figs. 3 a, b, and Liste des planches, p. 2.*
1848. Krauss. Stidafrik. Moll., p. 117 (mancinella).
1880. Von Martens. Mauritius G Seychellen, p. 235 (mancinella).
1892. Sowerby. Mar. Sh. S. Afr., p. 14 (mancinella).
1906. Hedley. Proc. Trans. Linn. Soc. N.S.W., xxxiil, p. 457.
1913. Smith. Proc. Mal. Soc., x, p. 288.
1919. Cooke. loc. cit., p. 92 (radula).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 170, pl. 7, fig. 6
(mancinella).
1956. Day & Morgans. Ann. Natal Mus., xiii, p. 307 (mancinella).
Living: Durban Bay (Cooke, S. Afr. Mus., and U.C.T.); Umpangazi
(Zululand) (U.C.T.); Delagoa Bay (U.W.).
Collected in Durban Bay by Burnup, and K.H.B. ‘Fairly common’ in the
harbour entrance (U.C.T.). The large specimen (v. infra: echinata) was taken
by the Pieter Faure at the jetty.
Distribution. Mauritius, Madagascar, Seychelles, Indo-Pacific.
Remarks. The large 6-whorled Pieter Faure specimen was returned by
Sowerby labelled as echinata (Blainv.). As it differs from the smaller specimens
only in having stronger tubercles I am assigning it to gemmulata.
In live shells (and in dried shells when wetted) the pale lirae on the apical
whorls make a pretty contrast with the reddish ground colour.
One specimen in S. Afr. Mus. (? H.M.S. Castor, east coast of Africa), 5
whorls, 42 X 31 mm., has an extra series of low blunt knobs below the suture
on 4th and 5th whorls, corresponding with the shoulder and peripheral knobs,
10 on 4th, 8 on 5th whorl.
Smith gave reasons for adopting Lamarck’s name instead of mancinella
Linn.
Thais (Cymia) carinifera (Lam.)
1822. Lamarck. Anim. sans Vert., vii, p. 241 (Purpura c.).
1836. Kiener. Cog. viv., p. 62, pl. 14, fig. 38 (Purpura c.).
1852. Souleyet. Voy. Bonite, ii, p. 603, pl. 40, figs. 4-6.
1911. Schepman. Siboga Exp. monogr., xlix, p. 354.
1919. Cooke. loc. cit., p. 93 (radula).
1952. Braga. Anais Est. zool. Ultramar., vii, 3, p. 77, pl. 3, fig. 6 (Cuma c.).
One somewhat worn specimen, 4 whorls, apex broken, 46 x 33 mm.,
dirty white, corresponding with Braga’s figure except that the lira above the
* In the S. Afr. Mus. copy, which belonged to Swainson, the name ‘mancinella’ is written
under fig. 3 in Swainson’s handwriting.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA AEF)
peripheral tubercles becomes crested at the outer lip and thus forms a
prominent tooth on the margin.
The specimen (S. Afr. Mus. A7688) is said to have come from Natal,
but more probably it came from Delagoa Bay, whence Braga recorded the
species.
Two young specimens, 22 (protoconch worn) X 15 mm., 4 postnatal
whorls, from Inhambane (U.C.T.) seem to belong to this species. They have
8 tubercles on each whorl (the above larger specimen has 9), 8-9 main lirae
between suture and peripheral tubercles, with intermediaries; numerous lirae
on base, the strongest one running from top of aperture and carrying tubercles
corresponding with the peripheral series; growth-lines crispate; costa on
rostrum rounded (not sharply keeled). Greyish, columella whitish, aperture
internally purplish-brown, operculum amber.
Radula with 180 and 220 rows (two radulae), central plate with median
cusp very long and narrow, 3 times as long as side cusps, latter with denticle
on inner side, externally 4-5 well-marked denticles, one or two of them
ascending the cusp (cf. Cooke’s description).
Distribution. Karachi, East Indies, Philippine Is.
Gen. Drupa Bolten
1919. Cooke. Proc. Mal. Soc., xiii, p. 100 (radulae).
This Indo-Pacific genus extends down the east coast of Africa to Natal.
The most southerly locality is Port Edward (living squamosa, U.C.T.). D. alfred-
ensis, although named after Port Alfred, probably came from Natal (if it was a
South African shell!). Sowerby’s record of tuberculata (granulata) from Port
Elizabeth is open to doubt; and Gould’s record of parvulum from Simon’s Bay
(False Bay) is certainly not acceptable (possibly Gould may have had a
Tritonalia scrobiculata or a young Thais dubia).
Drupa squamosa (Pease)
Fig. 50(a)
1868. Pease. Amer. 7. Conch., iii, p. 277, pl: 23, fig. 14 (Sistrum s.).
1919. Cooke. loc. cit., p. 101 (radula).
Radula with c. 190 rows, central plate with long narrow median cusp,
side cusp with one denticle on inner margin, externally 3—5 denticles, ascending
the cusp, lateral plate with base scarcely one-third width of central plate, blade
slender.
Living: Durban and Scottburgh (Cooke); Port Edward, Umhlanga, and
Kosi Bay (U.C.T.); Maxixe, Portuguese East Africa (U.C.T.); Mozambique
Island (U.W.).
Distribution. Indo-Pacific.
228 ANNALS OF THE SOUTH AFRICAN MUSEUM
Drupa anaxares (Duclos MS. Kiener)
1843. Kiener. Cog. vw. vil, p. 26, pl. 7, fie.17-
1919. Cooke. loc. cit., p. 105 (radula) (Morula a.).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 159, pl. 6, fig. 14
(anaxeres).
Living: Umkomaas, Natal (Cooke). Mozambique Island (U.W.).
Drupa granulata (Duclos)
Fig. 50(d)
1832. Duclos. Ann. Sci. Nat., xxvi, p. 111, pl. 2, fig. 9 (Purpura g.).
1832. Blainville. Nouv. Arch. Mus. Paris, i, p. 204, pl. 9, fig.
1919. Cooke. loc. cit., p. 106 (radula) (Morula g.).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 159, pl. 6, fig. 13
(tuberculata) .
Radula with c. 160 rows, central plate with strong median cusp, nearly
twice as long as side cusps, latter with a denticle on inner margin, externally
with one denticle, lateral plate with base about half width of central plate,
blade slender.
Living: Isipingo, Natal (Cooke); Durban (S. Afr. Mus., and U.C.T.);
Mozambique Island (U.W.).
Distribution. Indo-Pacific.
a) Sa
eA Pes
—_
Fic. 50.
Central and lateral radula plates of (a) Drupa squamosa (Pease); (6) D. granulata Duclos;
(c) D. (Cronia) margariticola Brod.; (d) Drupella sp. specimen C, with diagram of orientation of
lateral plates on the basal membrane; (e) Drupella sp. specimen B; ( f) Drupella sp. specimen A,
apex of lateral plate.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 229
Remarks. Cooke described a radula in the Gwatkin collection labelled
granulata from Isipingo, and a radula of a Mauritian tuberculata, in both of
which the median cusp of the central plate was shorter than the side cusps (this
might apply to the much-worn plates at the front end of a radula). The present
description is taken from specimens collected by myself at Durban, the shells of
which correspond with shells identified by Burnup.
Drupa squamiltrata (Smith)
Fig. 40(e)
1903. Smith. Proc. Mal. Soc., v, p. 377, pl. 15, fig. 17 (Szstrum s.).
Protoconch 2 whorls, alt. and diam. 0-75 mm., smooth. Postnatal whorls
5, profile angularly shouldered in middle of whorl or slightly below; axial
ribs 9 on Ist whorl, g-10 on following whorls, from suture to suture, broad,
the periphery an undulate outline in apical view, petering out on base;
crossed by one peripheral lira on 1st and 2nd whorls, by the peripheral lira
and one below it on 3rd, by 4 lirae above and one (or 2) below the peripheral
one on 4th and 5th whorls, 9-10 additional lirae on base, with fine inter-
mediaries. Growth-lines crispate. Costa on rostrum rounded. Columella
rimate anteriorly. Canal open; outer lip internally plicate. 13 x 7 mm.
Smith: 1g X 10 mm. White.
Operculum oval-reniform, nucleus in middle of outer margin.
Radula with c. 100 rows, central plate with median cusp much longer than
side cusps, latter with denticle on inner margin, but no denticles externally,
lateral plate rather slender.
Isipingo (Smith, coll. Burnup); Scottburgh (S. Afr. Mus., coll. Burnup).
Off Morewood Cove (near Umhloti, Natal), 27 fathoms, 4 alive; off
Umhlanga River, 22-26 fathoms, 3 (one alive) ; off Tongaat River, 36 fathoms,
faieedatment (5. Afr. Mus. PF. coll.). 29° 58'S. 31° 02’ E., 49 metres
eee. .).
Remarks. In general appearance like the Mediterranean and West African
Trophon fusulus (Brocchi) as figured by Knudsen (1956. Atlaniide Rep., 4, pl. 3,
figs. 4, 5).
Subgen. Cronia H. & A. Adams
Drupa margariticola Brod.
Fig. 50(c)
1795. Chemnitz. Conch. Cab., xi, p. 124, pl. 192, figs. 1851, 1852 (Murex undatus, part).
1832. Broderip. Proc. Zool. Soc. Lond., p. 177 (Murex m.). ~
1919. Cooke. loc. cit., p. 107 (radula) (Morula undata).
1938. Adam & Leloup. Mem. Mus. Roy. Hist. Nat. Belg., H.S. II, 19, p. 161, pl. 6, fig. 16.
Animal black. Radula with c. 135 rows, strongly ‘chitinized’, brown,
blackish posteriorly, central plate very wide, median cusp much stronger than
230 ANNALS OF THE SOUTH AFRICAN MUSEUM
side cusps, scarcely a trace of a denticle between them, lateral plate base wide,
more than half width of central plate, blade shorter than base.
Durban, Delagoa Bay (Sowerby); Isipingo, Natal (S. Afr. Mus. coll.
Burnup).
Living: Delagoa Bay, Nacala, and Mozambique Island (U.W.).
Distribution. Indo-Pacific.
Remarks. The radula of a Delagoa Bay specimen is quite different from
that described by Cooke from Karachi and Isipingo specimens, and resembles
that of D. (Cronia) amygdalus (Cooke, fig. 33).
Gen. DrRuUPELLA Thiele
1925. Thiele. D. Tiefsee Exp., xvi, p. 171, figs. 3, 4 (radula) (as subgen.).
1929. id. Handbuch, 1, p. 295, fig. 321 (radula) (as subgen.).
1939. Peile. Proc. Mal. Soc., xxiii, pp. 273-6.
Distinguished by the slender elongate lateral plates of the radula;
supposed to be an adaptation to feeding on certain kinds of coral (Cooke,
1895).
Drupella sp.
Fig. 50(@)-(f)
Three shells from Delagoa Bay (U.W.), 16-18 I0 mm., are so corroded
and overgrown with calcareous growth that identification is almost impossible
(cf. Peile). They seem, however, to agree with Scottburgh (Natal) examples in
S. Afr. Mus. collected and named by Burnup as elata Blainv. The outer lip in
all of them is internally denticulate, and the columella feebly granulate at the
anterior end. Two (A and B) have the columella and margin of outer lip white,
but farther inside the aperture is yellowish; the third (C) has the aperture
within, the columella and the outer lip margin pinkish-orange. Animals black.
Although externally the shells seem alike (except the coloured aperture
in C), their radulae differ in minor details. In general they resemble Peile’s
fig. 45 of a specimen from Samoa. Peile also recorded a second shell, with
similar radula, which had features ‘in common with ¢elata’.
Specimen A. At least 260 rows, central plate with very strong median
cusp, no denticle between it and side cusp, and no denticles or wrinkles exter-
nally, lateral plates 4-5 times as long as width of central plate, in ratio of
approximately 12 to 10 centrals (cf. Peile, p. 274), base not serrate, apex
bifid (fig. 50( f)).
Specimen B. About 100 rows, central plate with strong median cusp, no
denticle between it and side cusp, externally several wrinkles ending in fine
sharp denticles, lateral plates about 3 times as long as width of central plate,
in ratio of 2 or 3 to each central, base serrate, apex extremely finely bifid
(fig. 50(¢)).
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 231
Specimen C. About 150 rows, central plate with strong median cusp, a
tiny denticle between it and side cusp, the side cusp of the right side has also a
denticle about midway on its outer margin, externally no wrinkles or denticles,
lateral plates about 4 times as long as width of central plate, in ratio of 2 or
3 to each central, base not serrate, apex not bifid (not even on the unused
portion of radula) (fig. 50(d)).
In radula C it was observed that on the anterior third (c. 46 plates) the
laterals were directed forwards, on the hinder two-thirds (c. 104 plates) back-
wards. cf. Cancellaria (Barnard, 1958. 7. Conch., xxiv, p. 243, fig.) (fig. 50(d)
diagram).
Urosalpinx heptagonalis (Rve.)
Fig. 40(j)
1846. Reeve. Conch. Icon., iii, pl. 3, fig. 7 (Ricinula), and fig. 53 (Buccinum contractum).
1879. Smith. Proc. Zool. Soc. Lond., p. 201, pl. 20, fig. 32 (cnnotabilis).
1884. id. Zool. H.M.S. Alert, p. 47.
1903. id. Proc. Mal. Soc., v, p. 376 (contracta).
Ig1I. Schepman. Szboga Exp. monogr., xlix, p. 351 (contracta var. innotabilts).
Protoconch 2 whorls, alt. 0-5, diam. 0-6 mm., smooth. Postnatal whorls 6.
Axial ribs on 1st whorl 10-9, on 2nd and 3rd 9, on 4th and 5th 9-8, on 6th 8;
crossed by 2 spiral lirae on 1st and following whorls, on 3rd a fine lira between
them, increasing to 4 (5) on last whorl, fine spiral lirae also above the upper
and below the lower lira; complanate nodules at intersections with ribs; 15
additional lirae on base, of which the 5th (and sometimes also the 8th) is
stronger than the others, so that on the body-whorl there are 3 prominent
lirae: a peripheral pair, and one (sometimes 2) between these and end of
rostrum. Outer lip plicate within (at certain stages of growth). Growth-lines
scabrous. 33 X 18 mm.
Operculum oblong-oval, nucleus on outer margin.
Grey, drab, or fawn, the peripheral and the enlarged basal lirae usually
darker, or darker brown or bluish-grey where they cross the ribs; often also a
darker band below suture; aperture suffused within, brown at the margin;
operculum horny.
Radula with 90-100 rows, central plate wide, median cusp large, arising
from front margin of base, side cusp smaller, with a denticle between it and
median cusp, lateral plate rather slender; hinder rows coloured brown.
Natal (Krauss); Durban (Smith, and S. Afr. Mus.). Morewood Cove
(near Umhloti, Natal), 27 fathoms (S. Afr. Mus. P.F. coll.).
Living: off Umhloti River, 27 fathoms; off Tongaat, 36 fathoms (S. Afr.
Mus. P.F. coll.). Inyoni Rocks (Amanzimtoti, Natal), and 29° 58’ S.
30° 02’ E., 49 metres (U.C.T.). Delagoa Bay (U.W.). Inhambane (U.C.T.).
Mozambique Island (S. Afr. Mus. coll. K.H.B.).
Distribution. Indo-Pacific.
232 ANNALS OF THE SOUTH AFRICAN MUSEUM
Remarks. The upper whorls are frequently corroded or covered with
calcareous algal growth.
There are broad-shouldered and round-shouldered forms.
Smith (1879) remarked that Murex calcareus Dnkr. was very close to
innotabilis and Sowerby returned the P.F. specimen from Tongaat labelled
Tritonidea [sic] calcarea Dnkr.
Rapana bulbosa (Solander)
Fig. 51 (a)
1859. Chenu. Man. Conchyl., i, fig. 842.
1919. Cooke. Proc. Mal. Soc., xii, p. 102, fig. 29 (radula).
1942. Gravely. Bull. Madras Govt. Mus., n.s. V, 2, p. 48, fig. 8 a.
1952. Satyamurti. ibid., n.s. I, 2, pt. 6, p. 151, pl. 14, figs. 2 a, b.
Protoconch 2 whorls, alt. and diam. 1 mm., smooth but minutely pitted,
ending with a sharp plica. First postnatal whorl with numerous feeble axial
ribs, one spiral lira in middle of whorl and another below adjoining suture of
next whorl; both lirae become stronger on following whorls, especially the
upper one on which 8 or g vaulted squamae or hollow tubercles develop; the
number of these increases on later whorls, becoming solid tubercles or knobs.
100 (protoconch and 2 whorls missing) x 87 mm. (S. Afr. Mus.).
Radula (Cooke): central plate tricuspid, cusps broad, median one slightly
larger than side cusp, which has a denticle on its inner margin, externally
several wrinkles.
Durban (Smith, and (living) Cooke).
Off Umhloti River, 27 fathoms, 5 dead but fresh, 10-33 mm. long (S. Afr.
Mus. P.F. coll.).
Distribution. Indo-Pacific.
Gen AsPELLA Morch
1877. Morch. Malak. Bl., xxiv, p. 24.
1929. Thiele. Handbuch, i, p. 293.
Removed from the Cymatidae and placed by Thiele in the Muriczdae.
Radula formula 1.1.1, central plate tricuspid, a smaller cusp between the
median and side cusps, lateral plate unicuspid.
Aspella acuticostata (Turton)
Fig. 51(d)
1892. Sowerby. Mar. Sh. S. Afr., p. 9 (Ranella lamellosa, non Dnkr.).
1903. Smith. Proc. Mal. Soc., v, p. 378 (Ranella anceps, non Lam.).
1915. Bartsch. Bull. U.S. Nat. Mus., 91, p. 95 (anceps, non Lam.).
1932. Turton. Mar. Sh. Pt. Alfred, p. 109, pl. 24, no. 789 (Ranella a.).
Aperture (excl. canal) nearly twice in spire, canal 13 in aperture. Proto-
conch 14 whorls, alt. 0-5, diam. 0-6 mm., smooth, ending with a small plica or
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 233
varix extending from suture to suture, followed by a half whorl (perhaps this
is postnatal) with 3 plicae or varices the first of which ascends nearly to the top
of preceding whorl of protoconch. Postnatal whorls 5; each whorl with 6
varices, of which the end, 3rd, 5th and 6th are stronger than the others, and
extend farther over the suture on to the preceding whorl; a cross-section of the
shell is a parallelogram, not quite rectangular, with one of the prominent
varices at each corner; the strong varices extend across base of body-whorl to
rostrum. Growth-lines visible in some places; no spiral sculpture, but there are
faint indications of 6 or 7 crenulations on last varix (outer lip). 9 x 4 mm.
Creamy-white.
fe" pay
Mf
b
Protoconch of (a) Rapana bulbosa (Sol.).; (6) Aspella acuticostata Turton.
Port Elizabeth (Sowerby); Port Alfred (Bartsch, Turton); Umkomaas,
Natal (Smith); Still Bay (S. Afr. Mus. coll. Muir).
Remarks. The above description is taken from a specimen identified by
Tomlin as A. anceps Lam. Bartsch said the South African species was different;
he gave the type locality for anceps as west coast of America (presumably South
America, because it is not mentioned in Oldroyd, 1927. Stanford Univ. Publ.
Geol., II, pt. 2). If Bartsch is correct, Turton’s name must be accepted.
The specimen described is only slightly corroded, chiefly on the edges of
the varices. It is said to have come from Natal.
The other S. Afr. Mus. specimens, up to 11 (protoconch missing) x 4:5 mm.
are all beach-worn. From these the following characters are obtained: aperture
(excl. canal) 14 (juv. with 2 postnatal whorls) to 14 (3 whorls) to 2 times in
spire; canal 14 times in spire.
Turton gave the size as 15 X 6 mm.
In well-worn specimens the cross-section of the shell is nearly oval with
one prominent varix at each end (i.e. nos. 3 and 6), the other prominent
varices (2 and 5) being nearly obliterated. At first sight a worn and an unworn
shell appear rather different.
S. Afr. Mus. has 2 worn specimens said to have come from Mauritius
(Robillard coll.), the larger with 7 whorls (protoconch missing) 20 x 8mm. Each
whorl has 8 ribs or varices, i.e. 3 on each face between the main lateral varices.
234 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fam. COLUMBARUDAE
1922. Peile. Proc. Mal. Soc., xv, pp. 13, 14, fig. (radula) (near Muricidae).
1925. Thiele. D. Tiefsee Exp., xvii, p. 167 (in Muricidae).
1928. ‘Tomlin. Ann. S. Afr. Mus., xxv, p. 330 (separate family).
Characterized by the operculum: pear-shaped, tapering rapidly and
evenly to an acute nuclear apex, internal surface as in Fasciolaria; and by the
radula: central plate strongly concave in front, with 3 cusps, the middle one
the largest, lateral plate longer than its basal width.
Gen. COLUMBARIUM von Martens
Three forms occur at different depths off Cape Point, two of which are
regarded as new species. C’. radiale (Watson) and angulare n. sp. are both quite
distinct from rotundum n. sp.; radiale and angulare are not so distinct from one
another, but their habitats are separated by the area inhabited by rotundum.
Columbarium radiale (Watson)
Fig. 52(a), (4), (@)
1882. Watson. 7. Linn. Soc. Lond., xvi, p. 382 (Fusus r.).
1886. id. Challenger Rep., xv, p. 195, pl. 14, fig. 2 (Fusus r.).
1903. Von Martens. D. Tiefsee Exp., vii, p. 29 (Fusus r.).
Spiral lirae predominant. Aperture (incl. canal) 14-14 (or even slightly
more) times spire. Protoconch 24 whorls, alt. 2-3, diam. 2 mm. (but more or
less corroded), smooth, with indications of axial plicae on last half whorl.
Postnatal whorls 7, profile carinately angular. Axial ribs on 1st whorl 10-11,
increasing on following whorls to 13, and to 15—20 on last whorl, from suture
to suture on 1st and 2nd whorls, but thereafter evanescent above and below
the peripheral keel, on which they form bluntly triangular complanate lobes,
distinct as far as 6th whorl but feeble and indistinct on 7th; spiral lirae on Ist
whorl 3, i.e. one above and one below the prominent peripheral keel, on 2nd
and following whorls 2 above and 2 below, on later whorls there may be 2,
3 or 4 lirae above but only 2 (sometimes with a feeble intermediary) below the
periphery, and the lower one is often obscured by suture of following whorl;
15-20 additional lirae on base, widely spaced above, closer and less con-
spicuous on rostrum; close-set growth-lines between the lirae; suture incised,
more or less undulate; no parietal callus, columella straight or very slightly
curved; canal straight, narrow, distinctly marked off from, and 2-24 times
as long as rest of aperture, subequal to spire; outer lip not plicate within.
Periostracum thin, fibrous, fimbriate. Estimated length of largest specimen
(protoconch and tip of canal broken) 74 x 27 mm.; living examples
55 < 20 mim. Ay" 15 mam
Operculum as described under family, 9 x 6 mm. in 47 mm. shell.
White, periostracum buff or yellowish-brown, operculum amber-brown.
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 235
Radula with 120-130 rows, as described under family (fig. 52(e)).
Wead: 95° 4 8. 16°37 E., 150 fathoms (Watson); 34° 93’ S. 18° 21’ E.,
318 metres (von Martens). |
Living and dead: off Cape Point and west coast of Cape Peninsula,
90-230 fathoms (S. Afr. Mus. P.F. coll.).
Living: 35° 17'S. 18° 50’ E., 267 metres (s.s. Africana).
Remarks. Von Martens (1903) had already suspected this species to be a
Columbarium, and its appearance is so like a species of this genus that confir-
mation by the radula was no surprise.
Fic. 52.
(a), (b) Columbarium radiale (Watson), apex and later whorl. (c) C. angularen. sp. (d) C. rotundum
n.sp. (e) central and lateral radula plates of C. radiale (Watson).
Columbarium angulare n. sp.
Fig. 52(c)
Axial ribs distinct. Protoconch 24 whorls, alt. 1-5, diam. 1:3 mm., smooth
but a faint median lira on last half whorl. Postnatal whorls 7, profile angulate
but not carinate; axial ribs on 1st whorl 10-11, on 2nd 10-12, on 3rd and
following whorls 12 (sometimes 13), forming bluntly triangular lobes at inter-
sections with the peripheral lira, from suture to suture on early whorls, but
gradually receding from the suture above on later whorls, on body-whorl
distinct below periphery as far as level of top of aperture, evanescent on base;
spiral lirae distinct but not predominant, the peripheral lira well marked but
not carinate as in radiale, on 1st whorl 2 above and 2 below peripheral lira, on
_ and 3 above and 2 below, on grd 4 above and 2 below, increasing to (7) 8-9
above and 5-6 below, equal in strength but usually 2 of those below periphery
stronger than the others; on base at least 15-20 additional lirae with inter-
mediaries. Periostracum thin, fibrous, fimbriate. 51 (protoconch missing) x
13 mm.
Operculum and radula as in radiale. ‘The radula with 85-95 rows. Three
radulae were examined; in one of them the side cusps on the central plate were
36 ANNALS OF THE SOUTH AFRICAN MUSEUM
closely approximate to the median cusp, not separated by a V-shaped interval
as in the other two (and in the radulae of the other species).
White, periostracum pale buff, operculum amber-brown.
Off Cape Point, 720-900 fathoms, 9 specimens, 4 alive (S. Afr. Mus.
A4608 (Type)—A4611. P.F. coll.).
Remarks. A smaller and more slender species than radiale: 51 < 13 mm.
with 7 postnatal whorls whereas radiale has only 6 whorls at a size 55 X 21 mm.
Although the peripheral lira is strong, sometimes almost subcarinate, the
lobes at the intersections with the axial ribs are not complanate but 4-sided
pyramids, the ribs being more strongly developed than in radiale. The spiral
lirae above and below the periphery are more numerous.
Occurs in deep water beyond the slope of the continental shelf.
Columbarium rotundum n. sp.
Fig. 52(d)
Axial ribs predominant. Protoconch ? (corroded in all specimens). Post-
natal whorls 7, profile almost evenly convex with only a slight shoulder in the
middle of the whorl and slightly concave above. Apical whorls more or less
corroded, axial ribs on 3rd whorl 10, on 4th 10-11, on 5th 11-12, on 6th 12-13,
on 7th 13 (14), from suture to suture on 3rd whorl, but gradually receding
from suture above on later whorls, on body-whorl distinct below periphery as
far as level of top of aperture, evanescent on base; spiral lirae subordinate to
the ribs, on 3rd whorl 3 above, one at shoulder, 3 below, increasing on following
whorls to 7-8 above and 4-5 (unequal in strength) below; on base at least
15-20 additional lirae with intermediaries. Periostracum thin, fibrous,
fimbriate. 75 (protoconch missing) * 25 mm.
Operculum and radula as in radiale. The radula with 130-160 rows.
White, periostracum pale buff, operculum amber-brown.
Off Cape Point, 250-760 fathoms, numerous specimens living and dead
(S. Afr. Mus. A4592 (Type)—A4607. P.F. coll.).
Remarks. Distinct from radiale and angulare by the rounded non-angulate
profile of whorls, and the predominant axial ribs.
Seems to be subject to much more corrosion than either of the other two
species.
Occurs on the outer slope of the continental shelf in deeper water than
radiale, but not in such deep water as angulare.
Columbarium formosissimum ‘Tomlin
1928. Tomlin. Ann. S. Afr. Mus., xxv, p. 331, pl. 25, fig. 1.
Protoconch 2 whorls, bulbous, alt. 1-5, diam. 1-3 mm., smooth, junction
with rst postnatal whorl not sharply marked, the keel on the latter feeble at
CONTRIBUTIONS TO KNOWLEDGE OF S.A. MARINE MOLLUSCA 237
the start. Postnatal whorls 7 (incl. the Type, though Tomlin said 8). Peri-
pheral.keel on 1st and 2nd whorls not or only feebly undulate, on grd with 8
low and somewhat irregular undulations, on 4th with 8 well-marked but
slightly irregular undulations, on 5th with g lobes, on 6th 10, on 7th 11; the
lobes may be blunt or sharply triangular, sometimes hollowed in front. Perio-
stracum thin, fibrous, pale buff.
Radula with ¢. 100 rows.
Cape St. Blaize N. x E. } E., distant 65 miles, 85-90 fathoms; Cape Seal
N. x W. 4 W., 55 miles, 87 fathoms; Cape St. Francis NE. 29 miles, 75 fathoms;
Glendower Beacon (Port Alfred area) N. 4 W., 16 miles, 66 fathoms, one dead
fooeur. Mus. P.F. coll.).
Remarks. The Pieter Faure obtained 10 specimens. The Type and three
others (seen by Tomlin, now in S. Afr. Mus.) have their opercula, but Tomlin
made no mention of having extracted and examined a radula. Fortunately
there is one other shell containing the animal.
Columbarium natalense ‘Tomlin
1925. Tomlin. loc. cit., p. 331, pl. 25, fig. 2.
Protoconch 2 whorls, smooth, median keel faintly indicated on last half
whorl, alt. 1-3, diam. 1-25 mm. Postnatal whorls 64. Peripheral keel on Ist
whorl very feebly undulate, on 2nd 10 feeble undulations, on 3rd 11 distinct
lobes, on 4th the lobes beginning to be spiniform, on 5th and 6th whorls 12
more or less hollow spiniform, up-turned lobes.
Cape Natal (Durban) W. 2 N. distant 12 miles, 85 fathoms (S. Afr. Mus.
PP.’ coll.).
Remarks. Only one dead specimen was obtained, now in S. Afr. Museum.
The irregularity of the axial threads is due, not to abrasion as Tomlin’s
description might suggest, but to irregularities of growth; the shell is not at all
abraded, though the outer margin of the canal is broken just below the aperture.
As Tomlin remarked, distinctly resembling canaliculatum von Martens
from the Zanzibar channel, 400 metres.
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES
IN CAPE TOWN DOCKS
+ by
Ni Ac Be Minrarp, Prd).
Koology Department, University of Cape Town
(With 3 figures and 1 table in the text)
INTRODUCTION
Most of the material described in this paper was collected during a survey
of fouling organisms conducted in Table Bay harbour during the years 1946
to 1950, the results of which were published in 1952. During this period some
26 vessels were examined after dry-docking in Cape Town, and several series
of experimental plates were exposed for varying lengths of time in the water
of the harbour. In recent years a few additions have been made to the
collection as opportunity offered.
A study of the fouling community shows that the number of species which
settles on a floating structure is small in comparison with that in surrounding
waters. Thus, only 14 species of hydroids have occurred in the fouling as
against 65 species reported from False Bay near by (Millard 1957). Moreover
some species which are common on ships’ hulls and in harbour areas are
seldom encountered elsewhere (see also discussion). The fouling organisms,
in fact, make up a definite community of their own, the individuals of which
are particularly suited by their physiological or reproductive abilities to
occupy this ecological niche.
One surprising feature about the hydroid community is the fact that
some families, such as the Bougainvilliidae, Tubulariidae and Gampanu-
lariidae, are well represented, in numbers if not in species, and others, such as
the Sertulariidae and the statoplean Plumulariidae, are conspicuous by their
absence. The reason for this is not clear. Species which produce free medusae
and species with fixed gonophores are both represented and in approximately
equal numbers.
Grants from the staff research fund of the University of Gape Town made
possible the original work on fouling organisms and the purchase of micro-
scopic apparatus for the subsequent study of the material. The cost of publi-
cation was partly defrayed by a grant from the publications fund of the
University. Type material has been deposited in the South African Museum,
Cape Town (S.A.M.).
aoe
240 ANNALS OF THE SOUTH AFRICAN MUSEUM
List OF SPECIES
Tubulariidae Tubularia larynx Ellis & Solander
Tubularia warren Ewer
Corynidae Sarsia eximia (Allman)
Bougainvilliidae Bougainvillia macloviana (Lesson)
Bougainvillia ramosa (van Beneden)
Rhizorhagium navis n. sp.
Campanulariidae Laomedea angulata (Hincks)
Laomedea lovén Allman
Obelia bicuspidata Clarke
Obelia dichotoma (Linn.)
Obelia geniculata (Linn.)
Campanulinidae Lovenella chiquitita Millard
Plumulariidae Kirchenpaueria pinnata (Linn.)
Plumularia setacea (Ellis & Solander)
Family Tubulariidae
Tubularia larynx Ell. & Sol. 1786
Tubularia larynx. Allman 1872, p. 406, pl. 21. Hawes 1955, p. 333, figs. 1-5 (synonymy).
Material. One fruiting colony collected 24/5/47. Record number: SH 188.
Description. Stems branching, reaching a maximum height of 3 cm.,
annulated at intervals. Coenosarc with 2-4 internal longitudinal ridges of
endoderm, which may meet in the centre and divide the interior into canals.
The partitions themselves sometimes tubular.
Hydranth with 14-27 proximal and 15-19 distal tentacles. Distal
tentacles in a single verticil.
Blastostyles in 1 or 2 rows. Gonophores with 3-4 rounded tentacular
processes at distal end, less pronounced in the male than in the female.
Remarks. ‘This species has only once before been reported from South
Africa, from the Agulhas Bank by Stechow 1925.
Tubularia warrent Ewer 1953
Tubularia crocea. Millard 1952, p. 420, 428, 440, fig. 3.
T. warreni Ewer 1953; p. 351; figs. 1-4. Mullard 1959, p. 299.
Material. 10 colonies from ships’ hulls and 1 from experimental plates,
most of them very rich, and reaching a maximum height of 7-8 cm. Record
numbers: SH 2; 11) 195/95; 122; 176, 2545.95 7,304, 420", 420rK
Description. Stem with 2-5 (usually 2 or 3) internal longitudinal ridges of
endoderm, which when strongly developed may contain tubes. Mature
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 241
hydranth with 12-31 tentacles in proximal row, and 13-24 in distal row.
Largest hydranths reaching 6 mm. in length from distal tentacles to base of
dilation, and 2-5 mm. in basal diameter exclusive of tentacles.
Male gonophores usually without tentacular processes, but very occasion-
ally with 4-5 small conical processes. Female gonophores with 8 flattened
tentacular processes, of variable size. Actinula with 6-12 proximal tentacles
and up to 6 oral tentacles at liberation.
Nematocysts as described by Ewer.
Remarks. The importance of this species as a fouling organism was
discussed by Millard 1952 under the name of T. crocea. It is now known that
the colonies which occur so abundantly in the dock area are definitely Ewer’s
species as established by the examination of nematocysts in living specimens.
The early material brought in by ships was unfortunately not examined alive,
and the undischarged nematocysts of preserved specimens are of no help in
identification. No details of structure distinguish the latter from YT. warrenz,
and they have thus all been included with this species. Moreover, all the
ships concerned had had the opportunity of picking up the species in South
Africa at some time during their voyages.
The species is common on the hulls of ships and often forms a thick
carpet over the entire surface. It settles throughout the year, but mainly in
spring and summer. Ripe gonophores have been observed in all months from
January to August, and probably occur in the rest of the year too. Young
hydranths may bear gonophores at a height of about 1 cm., and about 35 days
after settlement.
As Ewer has suggested a study of the nematocysts in T. mesembryanthemum
may show it to be synonymous with T. warreni, and the former is well known
in Europe. It has not been recorded from South Africa.
T. crocea is distinguished from T. warreni only by its nematocysts and a
few minor features of the anatomy such as the absence of endodermal ridges
in the stem. This species has been reported from Liideritz Bay by Broch 1914,
and from a ship in the south Atlantic by Vanh6ffen 1910, who suggests that
the individuals may have settled in Simonstown (False Bay). It is possible
that in both these cases the material was in reality 7. warreni. (See also
discussion. )
Family Corynidae
Sarsia eximia (Allman) 1859
Syncoryne eximia. Allman 1872, p. 282, pl. 5. Hincks 1868, p. 50; pl. 9, fig. 2.
Material. 4 samples from ships’ hulls, among them one with numerous
young medusae, and another with 2 fruiting hydranths. Record numbers;
SH 1C, 123, 261B, 305.
Description. Stems tangled and richly branching, reaching a maximum
height of about 2 cm, Structure as described by Allman and Hincks, Hydranths
242 ANNALS OF THE SOUTH AFRICAN MUSEUM
bearing medusa buds at various stages of development, the oldest almost ready
for liberation but with the tentacles still unfurled. Medusa buds observed in
January.
Nematocysts: stenoteles, varying in size from 8-1 X 5:4 p to 15°3 X 9°9 L.
Remarks. This is the first sure record of the species from South Africa.
Non-fruiting material was reported by Day, Millard and Harrison 1952 from
Knysna Estuary as Syncoryne ?eximia, and by Millard 1957 from False Bay as
Coryne sp.
Family Bougainvilliidae
Bougainvillia macloviana (Lesson) 1836
Fig. 1, A-C
Perigonimus maclovianus. Vanhoffen 1910, p. 284, fig. 10.
Bougainvillia macloviana. Jaderholm 1923, p. 3. Browne and Kramp 1939, p. 284; pl. 14, fig. 6;
pl. 15, fig. 7-14.
Material. ‘Two small colonies, both fruiting. Record numbers: SH 196,
255B.
Description. Hydrorhiza forming a branching network over the surface of
barnacle shells and ascidian tests.
Stem unfascicled, slender, unbranched or branching irregularly several
times, narrower at base than at summit, reaching a maximum height of
3-4 mm. Perisarc roughly corrugated at base and on origin of branches,
smooth or wrinkled for the rest, continued over hydranth as far as the bases of
the tentacles as a ‘pseudohydrotheca’, covered throughout with adherent silt.
Hydranth with 8-12 tentacles in two close alternating verticils. Hypo-
stome conical.
Gonophores arising singly from the stems, branches or hydrorhiza, on
pedicels of variable length (usually shorter than gonophore), more or less
spherical when mature, completely enclosed in gelatinous perisarc. Oldest
medusa with 4 spherical marginal bulbs each bearing 2 black ocelli and 2
tentacles, a conical hypostome with a quadrangular base to its cavity, and 4
unbranched capitate oral tentacles.
Nematocysts of 2 types:
i. Undetermined heteronemes, 6:0-6:5 < 2:0-2°5 p.
ii. Desmonemes, 3°5-4°0 X 2°0-3°0 p.
Measurements (mm., preserved) SH 196 =SH 255B
Stem, diameter including perisarc a .. 0'05-O0°10 0°035-0°12
Pseudohydrotheca, height Ai a .. O*°13-0°23 0:*20-0°40
diameter sf ihe is sn .. 0°09-0°16 o-12-0°93
Gonophore (without perisarc), height
reaching ma a mF i si 0°24 0°30
diameter, reaching ., Vai i. HY 0:26 0°36
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 243
Remarks. There are certain slight differences between the two colonies,
though not sufficient to justify specific separation.
SH 255B is the sparser of the two, and the stem branches at most 2 or 3
times. The hydranths are slightly larger. The oldest medusa is ready for libera-
tion (fig. 1B); it has lost its connexion with the coenosarc of the pedicel and
has two of its tentacles unfurled.
SH 196 is a more luxurjant colony and is richly supplied with gonophores,
though these are not quite so advanced. Some have 4, and some 8, ocelli and
the marginal tentacles are still inturned. Oral tentacles are present in the
largest gonophores but could only be recognized in sections. Many empty
perisarcal capsules show that a crop of medusae has recently escaped.
The gonophores were observed in May and July.
The only previous record of the species from South Africa is that of
Jaderholm 1923.
Bian
Fic. 1. Bougainvillia macloviana (Lesson) (A—C), and Bougainvillia ramosa (van Beneden) (D-E).
A-C, 3 stems from SH 255B showing gonophores in various stages of development. D-E, two
portions of a colony from SH 429D showing filamentous appendages and young gonophores.
244 ANNALS OF THE SOUTH AFRICAN MUSEUM
Bougainvillia ramosa (van Beneden) 1844.
Fig. 1, D-E
Bougainvillia ramosa. Allman 1872, p. 311; pl. 9, figs. 5-7. Russell 1938, p. 152; 1953, p. 153,
fig. 74 A-C. Vervoort 1946, p. 135, figs. 52A, 53.
Bougainvillia vanbenedeni. Jaderholm 1909, p. 46; pl. 3, fig. 5.
Material. A bushy, well-preserved colony reaching a maximum height of
1°8 cm. (SH 429D), a single sterile stem of 0-8 mm. (SH 403), and a badly
preserved sterile colony with most of the hydranths disintegrated and reaching
ea Ged, (ielel ines).
Description. Stem unfascicled or weakly fascicled near base, increasing in
diameter from base to distal end, irregularly branched, or branches roughly
alternate in the distal regions. Perisarc smooth with very occasional groups of
corrugations on stem, corrugated or annulated on origins of branches, corru-
gated partly or wholly on hydranth pedicels, continued over hydranth to base of
tentacles as “‘pseudohydrotheca’. Filamentous appendages given off profusely
from stem, branches or hydranth pedicels, reaching a maximum length of
2°35 mm., occasionally branched.
Medusa buds scarce and present (in February) in the lower regions of
the colony only, mostly very young. Marginal bulbs visible, with stumps of
2 tentacles to each. Sections through the oldest stage show the beginnings of
4 oral tentacles.
Nematocysts of two types:
i. Microbasic euryteles, 8-0 < 3:0 p.
ii. Desmonemes, 4:0 X 2°5 p.
Measurements (ram., preserved, including perisarc)
Stem, unfascicled part, diameter .. ; he .. 0:08—0-20
Hydranth, approximate maximum Paneth a 22 SOS
maximum diameter .. ae ie Be a ores
Gonophore, maximum length we Be te ie heen
maximum diameter .. we bs a . sages
pedicel, length .. ‘G sit Fe Ar .. O'07-0'15
Remarks. ‘This material can be assigned to forma benedeni Bonnevie 1808,
which has been included in B. ramosa by most modern workers. It resembles
very closely Jaderholm’s figure (1909, pl. 3, fig. 5), except that the stem is
almost completely smooth.
B. ramosa has been reported from several localities on the south coast by
Stechow 1925.
Rhizorhagium navis n. sp.
Fig. 2
Holotype. SH Hea from the hull of a vessel which had not left Table
Bay, collected 10/2/58. (S.A.M, registered number H 124.)
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 245
Description. Colony creeping on weeds, other hydroids, etc. Hydrorhiza
giving rise to upright stems, each bearing a single terminal hydranth and
reaching a maximum height of about 5 mm. Hydrorhiza and stem covered
with perisarc, which terminates below the hydranth. Stem usually increasing
in diameter from the base upwards. Perisarc irregularly wrinkled or corru-
gated in parts, particularly near the base, often terminating in a swollen
rim. k
Hydranth with conical hypostome and 8-16 tentacles arranged in two
close, alternating verticils and held alternately elevated and depressed. Mouth
widely distensible and occasionally turned completely inside out.
Gonophores in the form of fixed sporosacs, borne on the stem below the
hydranth in an irregular fashion with the youngest above and the oldest below,
up to 8 per stem, male and female on separate colonies. Gonophore and its
pedicel covered with perisarc, which is thick below and very thin in the distal
region. Gonophores cryptomedusoid, without tentacle rudiments or radial
canals.
Male gonophores ovoid, with terminal opening, bearing the sexual products
around a swollen and hollow spadix.
Female gonophores ovoid, tapering below to short pedicel. Spadix swollen
and hollow, filling the gonophore in the proximal half, and bearing about
8 eggs, but often more, around the narrowed distal portion. Eggs developing
into planulae while still attached to the gonophore.
Nematocysts of 2 kinds:
i. Desmonemes, 3°5 X 2:0 yp.
ii. Microbasic euryteles, 6-5 xX 3:0 mw. Capsule elongated, slightly asym-
metrical, aperture off-centre. Butt about 2 length of capsule, thread coiled
transversely. Spines on butt not clearly determined. |
Colour: creamy white, with pink tinges in the hypostome of the hydranth
and spadix of the gonophore.
Development of gonophores. ‘This material was kept alive for two weeks in
the laboratory, during which time the development and maturation of the
gonophores was observed.
Gonophores of both sexes appear first as spherical hollow buds, covered by
ectoderm and endoderm (fig. 2B). Each bud lengthens and becomes pear-
shaped, and at the distal end, between ectoderm and endoderm, appears a
solid mass of cells, the entocodon (fig. 2G). As the gonophore continues to
elongate the proximal part of the endoderm remains closely applied to the
ectoderm so that the internal cavity of the spadix is spacious in this region,
but the distal part is narrow and separated from the ectoderm by the entocodon
which forms a sort of cap over it. The entocodon is now 2-layered, with a
shallow subumbrellar cavity between the layers, which is more evident in the
female than in the male. The inner layer of the entocodon is thicker than the
outer and in it the sex cells accumulate. A thin endodermal lamella is visible
346 ANNALS OF THE SOUTH AFRICAN MUSEUM
E
&
”
Fic. 2. Rhizorhagium navis n. sp.
A, part of a female colony drawn from living material. B—J, stages in the development of the
gonophores drawn from whole mounts. B, a young male gonophore. C and D, later stages in
development of male gonophore. E, the distal region of stage D on a larger scale to show details
of layers. F, a mature male gonophore. G, a young female gonophore. H, a mature female
gonophore. I, the female gonophore after the escape of the eggs, and J, with advanced planulae
still attached. ect, ectoderm. el, endoderm lamella. end, endoderm. ent, entocodon. g, germ
cells. fs, perisarcal covering. sc. subumbrella cavity.
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 247
between the entocodon and the ectoderm, but there are no radial canals or
tentacle rudiments (fig. 2D, E, G).
In the fully formed male gonophore the spermatogenic cells surround
the endodermal spadix for about two-thirds of the length, and this distal
region is sharply demarcated from the transparent proximal region (fig. 2F).
In the mature female gonophore the distal part of the spadix, surrounded
usually by a single tier of eggs, makes up about one-half of the length (fig. 2H).
Subsequently, either before or just after fertilization, the eggs escape from the
gonophore at the distal end, and with the release of pressure the spadix expands
to fill the whole space (fig. 21). The eggs remain attached to the gonophore
by a gelatinous perisarcal envelope and here develop into planulae which
reach an advanced stage before they finally escape (fig. 2J).
Measurements (mm., preserved)
Hydrorhiza, diameter #e a ea ae .. 0°08-0'14
Stem, diameter ee ae a oe .. 0°08-0-26
Hydranth, length from perisarc, normal position | gOr3O-1:29
maximum diameter, normal position : .. Of1Q—-0'53
Gonophore (without pedicel), male, length, eenehinie Fa) O02
G@iameter, reaching —.. ti ie - Sey Owl
Gonophore, female, length, reaching we ue a) 0:50
diameter, reaching .. + of Me sya MOPS
Remarks. ‘This material differs from most species of Rhizorhagium in that
the gonophores are borne on the stem instead of on the hydrorhiza. It resembles
most closely R. robustum (Warren) 1907 from Natal, but differs in the absence
of perisarc over the base of the hydranth, in the presence of more than one
gonophore per stem, and in the size and structure of the gonophore itself.
Family Campanulariidae
The classification of the Gampanulariidae is one of the most vexed
questions in hydroid systematics. Among modern workers there are two main
schools of thought: that of Broch (1918) who admits only two genera, Campanu-
laria and Laomedea (excluding Silicularia), and who is followed by most conti-
nental workers such as Kramp and Vervoort; and that of Stechow (1923c)
who recognizes as many as 17 genera.
Broch has distinguished his two genera mainly on the nature of the
diaphragm, and has discounted the method of reproduction, whether by fixed
sporosacs or medusae. His classification could not be accepted by medusa
systematists, and contributes little towards what should be our ultimate object,
a single composite classification for hydroids and their medusae.
While recognizing the fundamental importance of diaphragm structure,
the author also agrees with Rees (1957) that separate genera should be retained
for forms which produce free medusae and forms which produce fixed sporosacs.
248 ANNALS OF THE SOUTH AFRICAN MUSEUM
The difficulty then arises that various grades of degenerate or imperfectly
formed medusae occur in the family, and the grade may even differ in the two
sexes. It is proposed therefore for the purpose of classification to consider as
‘medusae’ only those which are fully formed and which can be classified by
the usual medusa keys, and as ‘sporosacs’ all imperfectly formed grades from
the styloid to the eumedusoid types. Thus we can retain Obelia for branching
forms with a true diaphragm and free medusae, and use Laomedea for branching
forms with a true diaphragm and fixed gonophores (including Gonothyraea,
Hartlaubella (= Obelaria) and Campalaria). Clytia can be retained for stolonial
forms (which may also branch in the form of a drepanium) which produce
free medusae. Stechow has shown that the diaphragm in the latter ranges
from the ‘Campanularia’ type to the ‘Laomedea’ type, and that in this respect
the genus is intermediate. The medusa is sufficiently different from that of
Obelia for generic separation. Campanularia can then be retained for forms with
an annular thecal thickening in place of a true diaphragm and which produce
fixed gonophores. As such it will include Orthopyxis with its degenerate medusae.
Laomedea angulata (Hincks) 1861
Laomedea calceolifera. Stechow 1925, p. 438.
L. angulata. Broch 1933, p. 100, fig. 43. Vervoort 1946, p. 305, figs. 134b, 135.
Campanularia calceolifera. Mullard 1952, p. 430.
Material. Two rich samples, both including male and female colonies
with abundant gonangia (borne in December and February). Maximum
height 2:3 cm. Record numbers: SH 327, 423.
Description. Structure and form of the colonies agreeing exactly with
previous descriptions. Stem unfascicled, branched or unbranched. No fila-
mentous appendages. Hydrothecae with slightly flaring margins.
Gonangia borne on the bases of the hydrothecal pedicels, usually
alternately on the anterior and posterior surfaces. Young ones truncated at
distal ends. Female gonangia containing a large number of eggs or fully
developed planula larvae. Male gonangia containing gonophores swollen
with spermatozoa.
Measurements (mm.)
Stem, diameter an i te i of .. O*10-0'15
Pedicel, diameter .. ah os i i .. 0°08—-0°13
Hydrotheca, height .. ee eg ne a .. 0°37-0°58
diameter at margin... i a * .. 0°24-0°42
height/diameter be % - at .) T=
Gonotheca, female, length .. De = tm) «42 OBS
diameter = a4 MY s a .. 0°39-0°57
Gonotheca, male, length .. a Mery ay Se .. 0°99-1°62
diameter : “ff a <3 th .. 0°29-0°45
Remarks. See discussion.
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 249
Laomedea lovén Allman 1859
Gonothyraea lovéni. Hincks 1868, p. 181; pl. 25, fig. 2. Allman 1871, p. 55, fig. 28.
G. loveni. Nutting 1915, p. 68; pl. 17, figs. 1-2. Millard 1952, p. 420, 440.
Laomedea loveni. Vervoort 1946, p. 310, fig. 137.
Material. 9 samples; rich colonies reaching a maximum height of 4-0 cm.
Record numbers: SH 4, 8, 124, 239, 255A, 261A, 271, 429C, 430B.
Description. Stems slender and branching.
Hydrothecae, and particularly the margins, extremely thin-walled and
delicate. Proportions variable, with length from 14 to over 24 times the
diameter. Marginal teeth with the typical truncated shape, separated by
rounded bays. No longitudinal striations.
Hydranth with 19-33 tentacles held alternately elevated and depressed,
completely retractable into hydrotheca.
Gonangia abundant, containing up to 8 gonophores, of which as many as
5 may be extruded as meconidia at one time. Female meconidia containing
about 5 planulae and bearing about 8 small tentacles. Male meconidia with
about 5 tentacles of the same length as the female. Gonangia observed in
January, February, and June to September.
Nematocysts of one kind only: basitrichous isorhizas, 6-7 XK 2°2 um.
Capsule elongated-oval, symmetrical.
Measurements (rm.)
Hydrotheca, length .. fe ey o, =. .. 0°39-0°63
diameter at margin... F i ge -- O-58-0-37
length/diameter oe - ue m: .. °59-2°73
Gonotheca, length, reaching a is ae ell S-C
diameter, reaching .. i 4 y: +, J0-36
Remarks. Due to the extreme delicacy of the hydrotheca the measurements
may not be wholly reliable, for in mounted specimens the margin is almost
invariably damaged, and the side walls tend to fall in, making the whole
structure appear narrower than it really is. For the same reason the base of
the hydrotheca often becomes telescoped on the pedicel making the diaphragm
appear oblique. In perfect, undistorted hydrothecae the length is usually
slightly over twice the diameter, and the diaphragm is perpendicular to the
hydrothecal axis. In end-on view there are very slight hollows on the outer
surfaces of the teeth, but these are far too shallow to give any impression of
striations in side view. This is the first record of the species from South Africa.
Obelia bicusfidata Clarke 1875
Laomedea bicuspidata. Vervoort 1946, p. 298, fig. 132; 1946a, p. 344, fig. 10.
Obelia bicuspidata. Millard 1958, p. 174.
Material. A small colony, reaching a maximum height of 6 mm., from a
floating dock from Calcutta. Collected 20/2/48. Record number: SH 341.
250 ANNALS OF THE SOUTH AFRICAN MUSEUM
Description. Stem unfascicled, unbranched or sparingly branched.
Hydrothecae rather small for the species and unstriated. The bicuspid
nature of the marginal teeth is not evident at first sight, for the bays between
members of a pair are practically equal in size to those between consecutive
pairs, but an end-on view of a hydrotheca shows the typical polyhedral outline,
with two internal keeled teeth arising from each plane.
Gonothecae present, but scarce.
Measurements (mm.)
Hydrotheca, length .. ue i i es .. O94-07%@
diameter hs Ae ae Ly eh: .. O16-oge
length/diameter ey Be ae i .. 1°59-1°89
Gonotheca, length .. oh Hf A. ae .. 0°52-0°62
diameter a A Me si =) Ori y ara
Remarks. This species has undoubtedly been transported from India,
whence it has been reported by Annandale 1915 as O. spinulosa.
Obelia dichotoma (Linn.) 1758
Obelia dichotoma. Millard 1952, p. 420, 426, 433, fig. 3; 1957, p. 198; 1958, p. 174.
Material. Numerous samples, some very rich, from ships’ hulls and
experimental plates. Maximum height 4:5 cm. Record numbers: SH 6, 7, 160,
220, 228, 2256, 279, 340, 349, 395, 398, 409, 410.
Remarks. This species has been discussed in a paper on fouling organisms
(Millard 1952). It settles mainly in the autumn, yet gonophores have been
observed in February, March, May to July, and September.
Obelia geniculata (Linn.) 1758
Obelia geniculata. Hincks 1868, p. 149; pl. 25, fig. 1. Millard 1952, p. 420, 4333 1957, p. 198.
Material. Two samples from experimental plates, reaching a maximum
height of 1:0 cm. Gonophores observed in March and June. Record numbers:
SH 347, 353-
Family Campanulinidae
Lovenella chiqutita Millard 1957
Fig. 3
Lovenella chiquitita Millard 1957, p. 198, fig. 7.
Material. A fairly rich colony growing on barnacle shells and other
hydroids. Record number: SH 430C (28/7/58).
Description. Pedicels arising directly from hydrorhiza or branching in a
sympodial manner as often as nine times. Colony reaching a maximum height
of 1°89 mm. Details of structure exactly as in original description.
HYDROZOA FROM SHIPS’) HULLS AND EXPERIMENTAL PLATES 251
Gonothecae plentiful, most of them empty, but two in the process of
discharging medusae (here described for the first time).
Medusa at time of liberation without apical process or peduncle, with 8
unbranched marginal tentacles of which the perradial ones are longer than
the interradial, without marginal or lateral cirri, with 8 closed adradial
marginal vesicles each containing two concretions, with a short stomach and a
simple quadrangular mouth, with narrow radial and circular canals, without
gonads, measuring approximately 0-3 mm. in depth and 0-4 mm. in diameter.
Colour: living hydranths colourless. Medusa transparent, with brown
patches on the bases of the tentacles and in the stomach.
Measurements. ‘The measurements are completely within range of those
quoted for the holotype and paratypes (Millard 1957), except for the gono-
thecae, some of which are slightly larger, the maximum size recorded being
0-70 mm. in length by 0-32 mm. in maximum diameter.
Remarks. Living material of this species was kept alive for a few days in
the laboratory. One medusa was in the process of escaping from its gonotheca
Fic. 3. Lovenella chiquitita Millard.
A-C from living material, and D from preserved material slightly shrunk. A, the young medusa
escaping from the gonotheca, B, ventral view of newly liberated medusa. C, the hydranth,
partially extended. D, lateral view of newly liberated medusa.
252 ANNALS OF THE SOUTH AFRICAN MUSEUM
at the time of collection, but died 2 days later without becoming completely
free. A second medusa successfully escaped 3 days after collection and was
fixed and mounted. |
The medusa, in the absence of lateral cirri, appears to belong to the
genus Phialella as defined by Rees 1939 and Russell 1953, rather than Lovenella.
These two genera, although closely related, are clearly distinguished by the
presence or absence of lateral cirri in the medusa, and by the nature of the
operculum in the hydroid, and are in fact usually placed in separate sub-
families. We are now confronted by a species in which the hydroid generation
belongs to one of these genera and the medusa to the other.
Kramp (1932a and earlier papers quoted therein) stresses the value of the
hydroid operculum in distinguishing sub-families of Campanulinidae, Lovenella
being included in the Calicellinae, and Phialella presumably in the GCampanu-
lininae, although the development of the operculum was described in only one
species of the latter, namely Campanulina lacerata (Johnston). The important
point is that the difference in the operculum in the two sub-families is the
result of a fundamental difference in development and should therefore be of
good systematic value.
In the present species the operculum clearly belongs to the ‘Lovenella’
type, and the only feature excluding it from this genus is the absence of lateral
cirri in the newly liberated medusa. Since it is possible that lateral cirri may
develop at a later stage it is proposed to retain the species in Lovenella, at least
until such time as the mature medusa is known and the development of the
operculum in Phialella is fully understood.
Family Plumulariidae
Kirchenpaueria pinnata (Linn.) 1758
Kirchenpaueria pinnata. Millard 1952, p. 420, 426, 433, fig. 3; 1957, p. 233.
Material. Abundant colonies from a variety of vessels and experimental
plates. Maximum height of colony 4:1 cm. Record numbers: SH 13, 14, 77,
145, 200, 245, 270, 272, 280, 282, 285, 288, 201, 202, 304, 313, B20 enema
361, 365, 368, 430D. |
Remarks. This species was discussed in a paper on fouling organisms
(Millard 1952). Gonophores have been observed in January, February, July
and October. Settling has been observed in most months of the year but
mainly in spring.
Plumularia setacea (Ell. & Sol.) 1755
Plumularia setacea. Hincks 1868, p. 296; pl. 66, fig. 1. Millard 1952, p. 420; 1957, p. 232; 1958,
p. 212. |
Material. Two small sterile colonies, with a maximum height of 0-7 cm.
Both colonies fall within forma typica Broch 1914. Record numbers SH 1B, 200.
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 253
DiIscussION
In all, 14 species of hydroids have been reported from floating structures
in Table Bay harbour, and an attempt was made to determine the origin of
the species, whether local or foreign (Table 1). Theoretically a history of the
vessels concerned (which had been fully recorded) should provide the answer,
but in actual fact it was no easy matter. Many of the vessels had had long
voyages with varied ports of‘call, often visiting Cape Town before their final
dry-docking, and sometimes lying idle in port for weeks or even months. Such
vessels might carry a mixture of foreign and local forms. Only one vessel (a
floating-dock towed from Calcutta) could be said to carry exclusively foreign
material, for she was dry-docked two days after arrival and had no time to
accumulate local forms. On the other hand one must bear in mind that
foreign species might be brought into the docks and reproduce there and even
establish themselves, eventually settling on vessels which have never left the
area.
Experimental) Purely | Purely | Local+- Total
local | foreign | foreign Distribution
vessels | vessels | vessels
Kirchenpaueria pinnata 5 - I 23 cosmopolitan
Obelia dichotoma 4 I 2 13 cosmopolitan
Tubularia warreni 5 - 5 II endemic to S. Africa
Laomedea lovéni 4 - 3 9 north Atlantic
Sarsia eximia 3 — I 4 cosmopolitan
Bougainvillia ramosa 2 = = 3 cosmopolitan
Plumularia setacea I - — 2 cosmopolitan
Obelia geniculata - - = 2 cosmopolitan
Bougainvillia macloviana 2 - _ 2 Antarctic
Laomedea angulata 2 - = 2 north Atlantic and
Falklands
Obelia bicuspidata I - cosmopolitan
Tubularia larynx = - cosmopolitan
endemic to S. Africa
endemic to S. Africa
Lovenella chiquitita
Rhizorhagium navis
TasB_eE 1. A list of the hydroid species in order of abundance, giving the number of records from
various sources. ‘Local’ vessels include those whose itinerary was restricted to the South African
coast.
Of the species listed, three, namely Tubularia warreni, Lovenella chiquitita
and the new species Rhizorhagium navis, are endemic to the country and could
only have a local origin. Yet the identity of 7. warreni is by no means finally
settled. The very fact that the species is characteristic of ships’ hulls and
harbour areas leads one to suspect that it is transported from place to place,
and possibly from Europe to South Africa, by ships, and to doubt whether it
should be held specifically distinct from such species as T. mesembyranthemum and
T. crocea which are abundant in harbours elsewhere. Only a full knowledge of
the variation of nematocysts in all related species can settle the problem,
254 ANNALS OF THE SOUTH AFRICAN MUSEUM
Of the remaining species all, with the exception of Obelia bicuspidata, have
been found on experimental plates or on vessels with a South African itinerary ;
some of the latter had never left Table Bay. Kirchenpaueria pinnata, Obelia
dichotoma, Plumularia setacea and Obelia geniculata in addition are common round
the South African coast, and can safely be assumed to have had a local origin.
Sarsia eximia, Bougainvillia ramosa and Tubularia larynx are by no means common,
but have all been recorded from the country on previous occasions.
Bougainvillia macloviana is an Antarctic form known to occur on ships’
hulls (Vanh6offen 1910). It can apparently penetrate as far north as South
Africa for it has been reported from off Borrocouto by Jaderholm 1923.
The records of Laomedea angulata and L. lovéni are interesting and sug-
gestive, for both are north Atlantic species and, apart from one record of
L. angulata from Simon’s Bay (Stechow 1925) and one from the Falklands
(Ritchie 1907a), they have apparently not been reported before from the
Southern hemisphere. They appear to be recent colonizers of the harbour area,
almost certainly transported there by ships from Europe or the Mediterranean.
Although well established, for they settle readily on newly exposed areas, they
have not spread to other parts of the coast. Stechow’s record of L. angulata
from Simon’s Bay was possibly also an immigrant from northern waters.
Only two species are of exclusively foreign origin, and were found on the
hull of the floating-dock from Calcutta mentioned above. These were Obelia
bicuspidata and O. dichotoma. Both are cosmopolitan in distribution and both
are also known from South Africa. O. dichotoma also occurs abundantly on
local vessels and experimental plates. These records provide supporting
evidence that species can be transported over long distances without serious
inconvenience, and that the genus Obelia is particularly hardy in this respect,
for the material was alive and reproducing freely on arrival.
In conclusion it is noteworthy that more than half (8 out of 14) of the
species recorded here are cosmopolitan in distribution, and 3 at least are on
the way to becoming so. One might pose the question: is their presence on
ships’ hulls a result of their world-wide abundance and ability to live under
varied conditions, or is their wide distribution a result of transportation by
ships ?
SUMMARY
A total of 14 species of hydroids is recorded. Amongst them is one new
species, Rhizorhagium navis, and one new record for the country, Laomedea
lovéni Allman. The medusa of Lovenella chiquitita Millard is described for the
first time. The composition of the hydroid fouling fauna is discussed and
suggestions made as to the origin of the species.
HYDROZOA FROM SHIPS’ HULLS AND EXPERIMENTAL PLATES 255
REFERENCES
Allman, G. J., 1871-2. A Monograph of the Gymnoblastic or Tubularian Hydroids. Parts 1
and 2. London.
Annandale, N., 1915. Fauna of the Chilka Lake. The Coelenterates of the Lake, with an
account of the Actiniaria of brackish water in the Gangetic Delta. Mem. Ind. Mus., 5,
65-114.
Broch, H., 1914. Hydrozoa benthonica, in Michaelsen’s Beitrige zur Kenntnis der Meeresfauna
Westafrikas, 1, 19-50. . 7
Broch, H., 1918. Hydroida, Part 2, in Danish Ingolf-Exped., 5, 1-205.
Broch, H., 1933. Zur Kenntnis der adriatischen Hydroidenfauna von Split. Norske Vidensk.-
Akad. Oslo, I. Mat.-Naturv. Klasse, no. 4, 1-115.
Browne, E. T. and Kramp, P. L., 1939. Hydromedusae from the Falkland Islands. Disc. Rep.,
18, 265-322.
Day, J. H., Millard, N. A. H. and Harrison, A. D., 1952. The Ecology of South African
Estuaries. Part 3. Knysna: A clear open Estuary. Trans. Roy. Soc. S. Afr., 33, 367-413.
Ewer, D. W., 1953. On anew Tubularian Hydroid from Natal. Ann. Natal Mus., 12, 351-357.
Hawes, F. B., 1955. Notes on the Variation occurring in Tubularia larynx Ellis & Solander.
F. Mar. Biol. Ass., 34, 333-346.
Hincks, T., 1868. A History of the British Hydroid Zoophytes, 1 and 2. London.
Jaderholm, E., 1909. Northern and Arctic Invertebrates in the Collection of the Swedish
State Museum (Riksmuseum). 4. Hydroiden. Kungl. Svenska Vetenskapsakad. Handl., 45,
I-I24.
Jaderholm, E., 1923. Hydroids from West and South Africa. Meddel. Géteborgs Mus. Zool., 26,
1-7.
Kramp, P. L., 1932a. The Godthaab Expedition 1928. Hydroids. Meddel. Gronland., 79, 1-86.
Millard, N., 1952. Observations and Experiments on Fouling Organisms in Table Bay Harbour,
South Africa. Trans. Roy. Soc. S. Afr., 33, 415-445.
Millard, N. A. H., 1957. The Hydrozoa of False Bay, South Africa. Ann. S. Afr. Mus., 43,
LS a ee
Millard, N. A. H., 1958. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part 1. Calyptoblastea. Ann. S. Afr. Mus., 44, 165-226.
Millard, N. A. H., 1959. Hydrozoa from the coasts of Natal and Portuguese East Africa.
Part 2. Gymnoblastea. Ann. S. Afr. Mus., 44, 297-313. -
Nutting, C. C., 1915. American Hydroids. Part 3. The Campanularidae and the Bonneviel-
lidae. Spec. Bull. Smithsonian Inst. Washington, 1-126.
Rees. W. J., 1939. A Revision of the Genus Campanulina van Beneden, 1847. Ann. Mag. Nat.
Fiist., ser. 11, 3, 433-447.
Rees, W. J., 1957. Evolutionary Trends in the Classification of Capitate Hydroids and Medusae.
Bull. Brit. Mus. (Nat. Hist.), Zool., 4, 455-534.
Ritchie, J., 1907a. The Hydroids of the Scottish National Antarctic Expedition. Trans. Roy.
Soc. Edinburgh, 45, 519-545.
Russell, F. S., 1938. On the Nematocysts of Hydromedusae. 7. Mar. Biol. Ass., 23, 145-165.
Russell, F. S., 1953. ‘The Medusae of the British Isles. Cambridge.
Stechow, E., 1923c. Zur Kenntnis der Hydroidenfauna des Mittelmeeres, Amerikas und
anderen Gebiete. Teil 2. ool. Jahrb. Fena, Syst., 47, 29-270.
Stechow, E., 1925. Hydroiden der Deutschen Tiefsee-Expedition. Wéissenschaftl. Ergebn.
Deutschen Tiefsee-Exped. 1898-1899, 17, 383-546.
Vanhoffen, E., 1910. Die Hydroiden der Deutschen Siidpolar-Expedition 1901-1903. Deutsche
Stidpolar-Exped., 11 Bd., Zool., 3, 269-340.
Vervoort, W., 1946. Fauna van Nederland. Aflevering XIV. Hydrozoa (Cl). A. Hydropolypen.
Leiden.
Vervoort, W., 1946a. Exotic Hydroids in the Collections of the Rijksmuseum van Natuurlijke
Historie and the Zoological Museum at Amsterdam. Zool. Meded., 26, 287-351.
Warren, E., 1907. On Parawrightia robusta gen. et sp. nov., a Hydroid from the Natal coast;
and also an Account of a supposed Schizophyte occurring in the Gonophores. Ann. Natal
Mus., 1, 187-208.
256 ANNALS OF THE SOUTH AFRICAN MUSEUM
ADDENDUM
The registered numbers of Hydroid type material previously described by N. A. H. Millard
and deposited in the South African Museum collections have not been published and are added
below. Numbers in parenthesis are University of Cape Town station numbers or serial numbers
of the collection made by the 8.8. Pieter Faure (PF).
Millard, N. A. H., 1955. “New species of Hydrozoa from South Africa’, Ann. S. Afr. Mus., 4% (5) 5
215-222.
Hydractinia altispina. Cotypes: S.A.M. H87 (F274), S.A.M. H88 (CP258), S.A.M. H89
(Bo2).
Hydractinia kaffraria. Cotypes: S.A.M. Hoo (BRE1r11A), S.A.M. Hot (HAM3Q). Para-
type: S.A.M. Ho2 (SUN3N).
Kygophylax cornucopia . Holotype: S.A.M. Ho3 (FB131B). Paratypes: S.A.M. Ho4 (TB1B),
S.A.M. Hg5 (FAL78S), S.A.M. Ho6 (FAL217N).
Millard, N. A. H., 1957. “The Hydrozoa of False Bay, South Africa’, Ann. S. Afr. Mus., 43 (4);
173-243-
Hydractinia canalifera. Holotype: S.A.M. Hg7 (CP332). ~~
Eudendrium deciduum. Holotype: S.A.M. Ho8 (FAL52V).
Halecium parvulum Bale, var. magnum. Holotype: S.A.M. Hog (FAL274R). Paratypes:
S.A.M. Hioo (FAL159L), S.A.M. H1i1 (PF405A), S.A.M. H30 (PF16287A).
Campanularia morgansi. Holotype: S.A.M. H2q4 (PF15675B). Paratypes: S.A.M. H7
(PF351C), S.A.M. Hior (FBrigL), S.A.M. Hroz (FAL26L), S.A.M. H32
(PF18232B).
Lovenella chiquitita. Holotype: S.A.M. Hio3 (FAL288J). Paratypes: S.A.M. Hro4
(FBr31F), S.A.M. Hro5 (FAL108.0).
Hebella furax. Holotype: S.A.M. H34 (PF18293B). Paratype: S.A.M. H106 (FALS58Y).
Synthecium hians. Holotype: S.A.M. H1o7 (FAL214G).
Sertularella capensis. Holotype: S.A.M. H1o8 (FBi14A). Paratypes: S.A.M. H1og
(FB115D), S.A.M. Hr1o (FAL64L).
Sertularella falsa. Holotype: S.A.M. H111 (FBi19C). Paratypes: S.A.M. Hi12 (FB131H),
S.A.M. H113 (CP333B).
Millard, N. A. H., 1958. ‘Hydrozoa from the coasts of Natal and Portuguese East Africa. Pt. I.
Calyptoblastea.’ Ann. S. Afr. Mus., 44 (5); 165-226.
Halecium inhacae. Holotype: S.A.M. H114 (IN140H).
Clytia serrata. Holotype: S.A.M. H115 (MOR2r16C).
Kygophylax geminocarpa. Holotype: S.A.M. H59 (PF12308A).
Kygophylax infundibulum. Holotype: S.A.M. H36 (PF10781B).
Hincksella corrugata. Holotype: S.A.M. H85 (PF12456J).
Sertularella dubia Billard var. magna. Holotype: S.A.M. H54 (PF12028B).
Sertularella mediterranea Hartlaub var. asymmetrica. Holotype: S.A.M. H116 (IN49K).
Sertularia linealis Warren var. longa. Holotype: S.A.M. H117 (IN140E).
Kirchenpaueria adhaerens. Holotype: S.A.M. H118 (RHB52G).
Monostaechas natalensis. Holotype: S.A.M. H79 (PF12456C). Paratypes: S.A.M. H48
(PF11803AF), S.A.M. H76 (PF12392G).
Monostaechas faurei. Holotype: S.A.M. H58 (PF12028F).
Plumularia irregularis. Holotype: S.A.M. Hi1g (DBN70Q).
Halicornaria africana. Holotype: S.A.M. H120 (AFR1028B).
| Halicornaria arcuata (Lamx.), var. epizootica. Holotype: S.A.M. H73 (PF12392D).
Thecocarpus giardi Billard, var. solidus. Holotype: S.A.M. H121 (AFR1028A).
16
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> en
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Out of print: Vols. I, II beck I-33 55 75 3), III (Ste ‘is bi r (Pate 4, 1,2
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLV
PART II, containing: —
Additions to the South African Museum Collection of Marine Fishes. By
F. H. Tarsot, M.Sc.
Note on Locality Records of Freshwater Fishes presented by F. D. McKean to
the South African Museum. By R. A. Juss.
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ADDITIONS TO THE SOUTH AFRICAN MUSEUM COLLECTIONS
OF MARINE FISHES
By
F. H. Tarsot, M.Sc.
ra . .
Barbourisia rufa Parr*
Parr, A. E., Barbourisidae, a new family of deep-sea fishes, Copeta, 1945, 127-129, 1 pl.
Two specimens, the locality of one ‘off Table Bay 1956’, no depth given,
and the other 32°10'S./16°15’E., 285 fathoms, 1959. (Registered numbers
S.A.M.19967 and S.A.M.22893 respectively.)
As the only record in literature of Barbourisia rufa is a single specimen
taken from the Gulf of Mexico in 1937, these two examples from the South
Atlantic off Cape Town are a most interesting find and greatly extend the
distribution of this species. The present two specimens differ slightly from the
180 mm. holotype, the main differences being: slightly smaller orbit, slightly
shorter jaws, and shorter gill rakers. In all other measurements and characteris-
tics they show very close similarity (see Table 1) except that in Parr’s description
of the genus he states ‘7 soft branchiostegal rays’, and the present specimens
have eight. As the thick skin covering the rays and their soft nature make
counting difficult it is probable that this will prove to be a miscount.
A pseudobranch of a few gill tufts is present. Lateral line pores between
6 and 16 shows the occasional raised pores mentioned by Parr. All pores are
small and without flaps, excepting the last which is very large, being the full
width of the lateral line tube.
Post-mortem colour is a brilliant scarlet-red (Ridgeway), which fades
rapidly on preservation to an off-white.
Pterothrissus belloct Gadenat
Cadenat, J. 1938. Note sur deux poissons nouveaux de la céte occidentale d’Afrique. Bull.
Mus. Hist. nat., Paris (2), 10, 361-369, 2 pl.
Poll, M. 1953. Poissons. III. Téléostéens Malacoptérygiens. Résult. sci. Expéd. océanogr. belge
Eaux cot. afr. Atlant. S., 4 (2), 1-258.
Four specimens trawled off Walvis Bay, 22°2’S./13°26’E., in 80 fathoms
(registered number $.A.M.21706). Donated by the Director, Division of
Fisheries, Department of Gommerce and Industries.
Poll (1953) has found this species to be abundant on the deeper portions
of the continental shelf off tropical West Africa. The present specimens are
the most southerly record.
*Since the above was written R. R. Rofen, Galathea Rep., 1, pp. 255-260, September 1959,
records and plates a specimen of B. rufa, from near Madagascar, mid-water, 450-700 metres.
This extends the range of the family to the Indian Ocean.
257
ITHSONIAN
Se STITUTION JUN 9
1953.
258 ANNALS OF THE SOUTH AFRICAN MUSEUM
Type S.A.M. S.A.M.
19967 22893
Standard length ai i a4: nf .. 180 mm. 297 mm. 294 mm.
Headii) ce 54 ye on ee Be 35°6 31 32
Snout .. ie ae a ee He ae Wiley Lion 11°6
Orbit diameter ap ts ay a Bi 4 pai! Py)
Upper jaw length a Se # ne af 25°6 22 23
Low jaw length che Ae he the ae 28 21 Ba
Greatest width of skull ae oP de oF 15 12°8 15
Interorbital distance .. =i Ay. KY Ge. 12" 10°4 LE=G
Snout to D. origin... Ns wie 3 st 64 61 61
Snout to A. origin... oe ae fs sas 69 68 71
Snout to V. origin... ne dik gs is 54-55 50 50
Dorsal fin base. . m7 a WY dibs aah 24 26 2
Anal fin base $A A Ene ue oa 18 17°5 16°3
Width of V. base a BS ae aah oe eae Ig 199
Widih of pectoral base es ats Ns aad ehab, Vee 2°4
Length of pectoral rays si as kes .. 6:°5-7 7° 4. 6-1
Length of ventral rays aie ane 1 .. 66°5 6-4 Bek
Longest dorsal ray .. A Mt, a es 1 8-4 8-2
Longest anal ray , sie dvs ail ee 10°5 8-4 9°5
Depth at shoulder sails we ae ae a 25 20 23
Least depth caudal peduncle oe oes ane 5 oon 8-2
D. to procumbent caudal rays ii oe bie 8 won| 8-2
D. to mid-base of caudal rays 6 a ~ 14-5 1471 15*3
Longest gill rakers .. se me a Bi Are 2°4 257
Dorsal count .. a se i aa as 20 20 20
Anal count a : ie ish ns 16 17 16
Number of lateral ee Sones set ia: Ss 29 29 30
Branchiostegal rays .. Bs st a a 7(?) 8 8
Gill rakers 5 me Lo Be ae . 64 14 6+ 15 6+ 15
TasLe I. A comparison of proportions of the type of Barbourisia rufa (from
Parr, 1945) with two South African specimens. Lengths are expressed
as a percentage of standard length.
Allothunnus fallai Serventy
Serventy, D. L. 1948. Allothunnus fallai, a new genus and species of tuna from New Zealand.
Rec. Canterbury (N.&.) Mus., 5, 131-135.
One specimen, fork length 835 mm. taken by spear gun in 3 fathoms off
Millers Point, Cape Peninsula, on 8 April 1958, and donated by Mr. D.
Hammond (registered number S.A.M.21546).
The Slender Tunny, with its reduced dentition and high gill-raker count,
is stated by R. A. Falla (in a note appended to Parrot, 1958, Rec. Dom. Mus.,
Wellington 3, p. 119) to be ‘not uncommon in southern New Zealand seas, but
rarely caught’. Serventy’s original 3 specimens were from South Island, New
Zealand, south of 43°, and Falla mentions sight and photographic records from
the Auckland Islands (50° S.), so it can presumably be considered a cold-water
tunny. This is the first South African record.
The present specimen shows some small differences from the original
description (see Table II), but there seems no doubt that it is conspecific.
ADDITIONS TO S.A. MUSEUM COLLECTION OF MARINE FISHES 259
Scaling is not as complete as in the type. Behind the distinct corselet of larger
scales the body is covered in fine scales on its upper half to below the lateral
line and the lower half of the sides and the belly are naked. The vomer and
palatines are slightly rough to the touch, being covered in microscopic granular
teeth. Serventy doubtfully gives a vertebral count of 41. X-ray photographs
of the present specimen show 39 vertebrae however.
Fin counts: Dorsal XVII 12 plus 7; anal 13 plus 7. Gillrakers 23 plus 48
left, and 22 plus 53 right. *
The specimen is a mature male.
Post-mortem colour was steely blue above shading to silver below, and
with no distinctive markings. A photograph taken immediately after death
shows a dark line from the pectoral tip, running longitudinally and upwards
to the 3rd dorsal finlet.
Type S.A
Fork Length (snout to caudal fork) a ah ae .. 616 mm. 850 mm.
Diameter of eye - a Life a sub We is Q°7 3°6
Head Length .. of; Bee aie Se ef, = sie 26 26
Snout to origin of pectoral .. me Sp yp te as 27 28
Snout to origin of first dorsal or MA st se ba 31 30
Snout to origin of second dorsal .. ss a st ae 63 59
Snout to origin of ventral .. st An ee a ce 28 31
Snout to vent .. : 64 61
Depth of body at Sees of Pa D. te = Speck greatest t depth) a1 20
Depth of body at vent ae ‘ ; 18-8 17°4
Length of pectoral .. : aif Ae oi és 10°5 11-2
Length of pectoral pared. pode aN Semen cre a 12°2 12-2
Inter-orbital distance .. ae Be sy o Ae ea 7°8 a2
Length of maxillary .. : : ae me wf bie 9°3 9°3
Snout to hinder edge of gece is sts ae ate 20°1 19°8
Height of first dorsal . oY Ke am oy oie ay 10°4 8-1
Height of second Heed 7°9 7°5
Height of anal 5 ie ae 7°8 7°78
Snout to anterior nostril ae Sue ee is ms ‘e 4°6 4°9
Snout to posterior nostril wg aS af be An 6-8 6-9
Longest gill raker ue ee be ue ue ie ie 4°5 452
Tas_e IT. A comparison of the proportions of the type of Allothunnus fallai
with the South African specimen. Fork length is in mm. and all other
measurements are expressed as a percentage of fork length.
~NOTE ON LOCALITY RECORDS OF FRESHWATER FISHES
PRESENTED BY F. D. McKEAN TO THE SOUTH AFRICAN MUSEUM
By
R. A. Juss
Department of Ichthyology, Rhodes University, Grahamstown
Gilchrist and Thompson (1917) have recorded species of fishes presented
by Mr. F. D. McKean and have given the locality as Sawmills, Bulawayo,
Rhodesia. Sawmills is actually a railway station approximately 60 miles north
of Bulawayo, on the Bulawayo-Victoria Falls railway line, and situated on
the Umgusa River. The Umgusa is a tributary of the Gwaai River which
belongs to the Middle Zambezi River system. The fish fauna of the Upper
Zambezi system above the Victoria Falls differs from that of the Middle
Zambezi system (Jubb, 1958, p. 178). Except for Tilapia melanopleura, which is
widely distributed, the species represented in McKean’s collection are found
only in the Upper Zambezi system and the Kafue River, and not in the Middle
Zambezi River system; as a typical example the predatory Serranochromis,
much sought after by anglers, are conspicuous by their absence from this
section of the Zambezi River. It would appear that Mr. McKean’s address,
prior to 1917, has been recorded as the locality from which the specimens
were collected. It has not been possible to get any information about
Mr. McKean.
The names listed are in accordance with Barnard’s (1949) revision of
the South African Cichlidae. I have personally examined the Serranochromis
specimens in the South African Museum which were presented by Mr. McKean.
It is highly probable that these fishes actually were collected in the Zambezi
above the Victoria Falls.
. 548 Ctenopoma multispinis Peters.
. 538 Recorded as Pelmatochromis robustus, but is Haplochromis smith Castelnau.
. 526 Three specimens recorded as Serranochromis thumbergi Castelnau.
. 525 Two specimens recorded as Serranochromis angusticeps Boulenger.
499 One specimen Tilapia mackeani Gilchrist & Thompson, 1917, which
is a synonym of Tilapia melanopleura Dumeril.
p. 482 Two specimens Tilapia intermedia Gilchrist & Thompson, 1917, which
are synonyms of Tilapia macrochir Boulenger.
p- 481 Tilapia kafuensis Boulenger.
ode oo
BARNARD, K. H. 1949. Revision of South African Cichlidae. Rep. Inland Fish. Dept., C.G.H.,
No. 5 (1948), 48-61.
Giucurist, J. D. F. & THompson, W. W. 1917. The freshwater fishes of South Africa. Ann.
S. Afr. Mus., 115 465-575.
Juss, R. A. 1958. A preliminary report on the collections of freshwater fishes made by the
Bernard Carp expeditions to the Caprivi Strip, 1949, the lower Sabi River, 1950, and
to Barotseland, 1952. Occ. Pap. nat. Mus. S. Rhod., No. 22B, 177-189.
260
ANNALS
SOUTH AFRICAN MUSEUM
VOLUME XLV
PART III, containing :—
The Polychaet Fauna of South Africa. Part 5. Errant Species dredged off Cape
Coasts. By J. H. Day. (With 14 figures in the text.)
(a din T ea vin
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ISSUED MAY 1960 ~~ PRICE 16s.
PRINTED FOR THE
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vrs
THE POLYCHAET FAUNA OF SOUTH AFRICA
PART 5. ERRANT SPECIES DREDGED OFF CAPE COASTS
by
J. H. Day
Professor of Zoology, University of Cape Town
CONTENTS
page page
Introduction .. “2 as = 201 Systematic account ... ae soe 27K!
List of collecting stations .. ne) 204), Weferences ot a i ila ie af
INTRODUCTION
Earlier papers in this series have dealt with the estuaries and intertidal
fauna of Mogambique, Natal and Cape Coasts. This is the first account of
species which occur below tide-marks and it will be evident from the long lists
of collecting stations that a great deal of field-work has been done since the
work first started in 1947. Actually the prime object of this dredging survey is
to determine distribution patterns around the southern part of Africa and not
only the Polychaeta but the whole benthonic fauna is being surveyed. Many
systematic reports on the other groups have been published and further work
is in progress. All the records are being entered on card catalogues and when
the area has been adequately covered a biogeographic analysis will be attempted.
Meanwhile many new species are being discovered.
The material has been obtained from many sources. I am indebted to the
Director of the Division of Fisheries for dredgings brought in by R.S. Africana II
and Palinurus and I also wish to thank Messrs. Irvin and Johnson for allowing
students to collect specimens on board commercial trawlers. The Hydrographic
Section of the South African Navy generously provided facilities for dredging
on board S.A.S. Natal during three cruises between Port Elizabeth and Durban
and Dr. Nafe of the Lamont Geological Observatory kindly allowed me
facilities during the cruise of the Vema between Cape Town and Durban. To
all of these organizations I tender my thanks but the bulk of the material has
been collected by members of my own department working on small boats
_ during university vacations. In this way rich hauls were made in Lamberts
Bay, Saldanha Bay, Table Bay, False Bay, Mossel Bay and Algoa Bay. The
expenses of such trips were covered partly by grants from the Staff Research
Fund of the University and partly by grants from the South African Council
of Scientific and Industrial Research. The latter organization has also paid the
261
SMITHSONIAN .
INSTITUTION JUN 9S 1868
262 ANNALS OF THE SOUTH AFRICAN MUSEUM
salary of my research assistant for many years and Dr. John Croil Morgans
made some very valuable collections by diving in False Bay during his tenure
of the post. A full report of his diving survey has recently been published
elsewhere (Morgans 1959).
Earlier records of errant polychaets dredged off the Cape coasts will be
found in McIntosh (1885 and 1904), Ehlers (1908 and 1913), Ramsay (1914),
McIntosh (1925), Monro (1930 and 1936), Day (1934) and Treadwell (1943).
The species recorded by Augener (1918 and 1931) from dredgings off South
West Africa must be included for there can be no doubt that this area has a
similar fauna to the Western Cape.
Altogether these earlier workers recorded 74 species of errant polychaets
from below tide-marks. The present paper contains new records, notes or full
descriptions of 171 species. 61 of them are species known from the earlier
dredgings, 42 are species previously known only from the shore, 34 species
are new records for South Africa, 22 are new species and 12 are doubtful
species or new varieties. The full total of species now known from dredgings
around the Cape or South West Africa is 184 and for the sake of convenience,
the 13 species recorded by earlier workers and not included in the systematic
section of this paper is given below.
Aphroditidae
Eunoe assimilis McI.—MclIntosh 1925
Eunoe macrophthalma Mc.1.— McIntosh 1925
Lagisca hubrechtt Mcl.—Monro 1930
Macellicephala mirabilis Mc1.—MclIntosh 1905, 1925
Polynoe caput-leonis Mc1.— McIntosh 1925
Panthalis oerstedi var. capensis Mc1.—MclIntosh 1925
Leama hystricis Ehl.— McIntosh 1925
Hesionidae
Magalia (=Syllidia) capensis (McI.) —MclIntosh 1925
?Irmula spissipes Ehl.—Augener 1918
Syllidae
Sphaerosyllis perspicax Ehl.—Augener 1918
Nereidae
Nereis pelagica Linn.— Ramsay 1914
Eunicidae
Eunice grubei Grav.—Ehlers 1908a
Onuphis quadricuspis Sars— McIntosh 1925
THE POLYCHAET FAUNA OF SOUTH AFRICA 263
The other names which occur in the literature are synonyms of species
described in this paper. Many of them are misidentifications of European
species and one of my main tasks has been to eliminate these names from the
South African faunistic lists. Unfortunately the descriptions are often incomplete
and it has been necessary to examine the original material. During 1952 the
C.S.1I.R. provided me with funds to visit the British Museum and examine the
South African material housed there and to compare my own collections with
the types.
I wish to thank the Director of the British Museum and Mr. Norman
Tebble of the Annelid Section for their kindness and help on this occasion
and during a subsequent visit in 1958. I also wish to thank the Directors of
the Scottish National Museum, the Swedish State Museum, the Hamburg
Museum, the Berlin Museum and the U.S. National Museum for sending me
South African specimens lodged in their respective institutions.
One of the most important results of this sort of work was a general
review of the genus Diopatra, which has been reported, not only from South
Africa, but also from many other parts of the world under the name Diopatra
neapolitana. An examination of material from the type locality (Naples) showed
that the great majority of the records are misidentifications. A general discussion
of Diopatra will be found on p. 338; the point which is stressed here is that
similar work on difficult genera such as Eulalia, Exogone, Autolytus, and
Lumbrineris suggests that a re-examination of type material or, where this is
lacking, of material from the type locality is well worth while. It will lead to
the solution of many anomalies of distribution and it now seems very probable
that species of Polychaeta are by no means as widespread as has been supposed.
Distribution patterns in this group will probably be found to follow the lines
which Ekman (1953) has proposed for the bulk of the marine fauna.
There is no doubt that many new species of errant polychaets await
discovery in Cape waters. The University now has its own research vessel
and almost every dredging brings up species new to the area or new to science
and the deeper waters off the Eastern Cape have hardly been explored. For
this reason no systematic key is included in this paper although one has been
produced and is constantly being revised. Further work on errant species must
wait until the bulk of the sedentary species has been described. This will form
the subject of the next paper in this series.
STATION Lists
I must apologize for not giving the full station data below each species.
It is realized that this causes a certain amount of inconvenience but space
does not permit the full collection data to be repeated in this way. Full details
for each dredging or diving station are given below, and under each species
will be found only the station number with the number of specimens in brackets,
e.g. AFR.728(1) under Aphrodite alta means that 1 specimen was obtained by
R.S. Africana II and reference to the station list will give full details of date,
264 ANNALS OF THE SOUTH AFRICAN MUSEUM
position, depth and type of bottom in the conventional abbreviated form.
The sequence of the station lists is from the west or Atlantic coasts around the
Cape towards the Eastern Province and Natal although some of the Africana
(AFR) and Trawler (TRA) stations cover a wide range of coastline. In some
cases the number of specimens obtained at a particular station was not
accurately counted but was noted as common indicated as (c), fairly common
(fc), or merely present (p). These letters in brackets are thus shown against the
relevant dredging stations.
The types described in this paper will be deposited in the South African
Museum, Cape Town.
The Trustees of the Museum gratefully acknowledge the grant in aid of
publication of this paper received from the South African Council for Scientific
and Industrial Research.
STATION DATA
LAMBERT’S BAY Drepcinc (LAM)
No. Date Position Depth Bottom
(Metres)
LAM. 1 16.1.57 32.04.39/18.18.2E 15 Ss.
LAM. 4 do. do. do. do.
LAM. 5 L757 32.04.59/18.18.E 17 Sh.R.
LAM. 6 17.1.57 32.04.79/18.18.5E 9 Serie
LAM. 8 18.1.57 32.059/18.17.9E 23 S. Sh. R
LAM. to 17.1.57 32.04.79/18.17.7E 23 S. Sh.
LAM. 11 18.1.57 32.059/18.17.7E 29 S. Sh.
LAM. 13 19.1.57 32.049/18.18.1E 18 R.
LAM. 15 18.1.57 32.059/18.17.7E 17 S. Sh. R
LAM. 16 Iigoliay 32.04.85/18.18.2E II Ss.
LAM. 17 afta a) 32.05.39/18.17.4E 23 S.
LAM. 18 18.1.57 32.04.859/18.17.8E 17 R:
LAM. 19 do. do. do. do.
LAM. 22 17.1.57 32.07.59/18.17.6E 20 S. R.
LAM. 23 17.1.57 32.04.15/18.18.6E 15 S. Sh.
LAM. 24 16.1.57 32.04.68/18.18.2E 17 R.
LAM. 25 1 seis 32.04.25/18.18.4E 8 S. Sh. R.
LAM. 26 18.1.57 32.04.99/18.17.5E 27 S. Sh. R
LAM. 27 16.1.57 32.04.19/18.18.4E 16 R.
LAM. 31 19.1.57 32.05.19/18.17.7E 20 R.
LAM. 33 19.1.57 32.05.29/18.17.5E Plankton sample.
LAM. 35 19.1.57 32.05.59/18.17.7E 27°5 Sh. R.
LAM. 38 19.1.57 32.05.49/18.17.7E 27 S. sh:
LAM. 39 19.1.57 32.05.49/18.17.6E 30 S. Sh.
LAM. 40 19.1.57 32.05.59/18.17.6E 28 S. Sh.
LAM. 41 21.1.5 7 32.059/18.17.7E 20 S. Sh.
LAM. 43 21.1.57 32.04.99/18.18.2E 13°5 Sis
LAM. 44 21.1.57 32.04.75/18.17.6E 20 R.
LAM. 45 21.1.57 32.059/18.18.2E 8 DB. Ike
LAM. 47 21.1.57 32.04.49/18.17.7E 23 R.
LAM. 48 22.13.57 32.045/18.17.9E B27 S. Sh.
LAM. 49 21.1.57 32.04.85/18.18.1E 10 S. R.
LAM. 51 23.1.5 7 32.08.59/18.17.7E 16°5 ae
LAMBERT’s Bay Drepcinc (LAM)
No. -
Date
21.1.57
23.1.57
22.1.57
23.1.57
23.1.57
23-1.57
23.1.57
23.1.57
23.1.57
23.1.57
19.1.57
Position
32.04.79/18.18.2E
32.099/18.17.8E
32.04.25/18.17.7E
32.115/18.18.1E
32.109/18.18.1E
32.095/18.18E
32,02S/18.18E
32.12S/18.17.9E
32.01.59/18.18E
32.01.59/18.17.8E
32.05.59/18.17.7E
SALDANHA Bay Drepcinc (SB)
No.
SB.113
SB.114
SB.115
SB.116
SB.117
SB.118
SB.119
SB.120
SB.121
SB.122
SB.125
SB.127
SB.129
SB.130
SB.132
SB.135
SB.136
SB.143
SB.144
SB.145
SB.173
SB.175
SB.177
SB.179
SB.180
SB.181
SB.183
SB.184
SB.189
SB.193
SB.195
SB.197
SB.199
SB.202
SB.203
SB.205
SB.207
SB.208
Date
Position
33.00.75/17.59.8E
33.00.459/17.57.5E
do.
33.00.1S/17.59.2E
33.00.39/17.58.5E
33.01.59/17.58E
33.02.85/18.01.2E
33.03.45/18.01.8E
33.059/18.01.4E
33.04.95/18.00.4E
33.01.49/17.57-7E
33.04.6S/17.59.8E
33.04.59/18.00E
33.04.6S/18.00.6E
33-045/17.59.3E
33.035/17.58.6E
33.035/18.00.5E
33.05.15/18.01.2E
33.05.39/18.01E
33.04.85/18.00.5E
33.05.19/18.01.5E
33.02.85/18.00.6E
33.035/18.00.gE
33.03.6S/18.00.4E
33.03.59/17.58.5E
33.01.65/17.59.3E
33.02.59/17.58.7E
33.01.59/17.58.8E
33.01.19/18.00.3E
33.00.79/17.58.4E
33.03.59/17.59.2E
33.04.45/17.50.4E
33.01.79/18.01.4E
33-03-59/17.57-5E
33-05.59/17.55-5E
33.03.6S/17.56.4E
33.02.59/17.57.5E
33.01.99/17.56.3E
Dept
(Metres)
Depth
(Metres)
~
[o)
Qu
La]
OND HUNTOW OANA OM
-_
m ND
Oe WN
on
THE POLYCHAET FAUNA OF SOUTH AFRICA
- Bottom
DDD win
N
in
Bie
Bottom
wn
do.
APnnn
cn
re)
DnnAY
Ss
265
266
ANNALS OF THE SOUTH AFRICAN MUSEUM
LANGEBAAN LAGOON DREDGING (LB)
No.
LB.155
LB.158
LB.159
LB.160
LB.161
LB.169
LB.190
LB.191
LB.239
LB.299
LB.300
LB.323
LB.363
LB.364
LB.38o0
LB.382
LB.391
LB.456
LB.472
LB.496
Date
15.7.46
do.
do.
TasLe Bay Drepcinc (TB)
No.
TB.3o1
TB.302
TB.303
TB.304
TB.305
TB.306
TB.307
TB.308
TB.309
TB.310
TB.312
TB.313
TB.314
Date
4.8.46
11.2.4.7
do.
15.12.57
TB.315-333 do.
Position Depth Bottom
(Metres)
33.07.15/18.02.4E 2 f. S.
33.099/18.04.2E 4 do.
33.105/18.04.8E 4°5 f. S. M.
33.06.48/18.01.9E 3 S. Sh.
33.05.6S/18.00.8E 5 Sh. R.
33.10.59/18.03.8E 2 i. 0.
33.11.39/18.05.5E 0-2 f. S. M.
do. do. do.
33.079/18.02.7E 2 f. S.
33.06.85/18.01E 2°5 S. Sh.
33.07.69/18.02.3E a Ss.
33.06.8S/18.01E 2°5 S. Sh
33.07.15/18.02.7E 4 S
33.05.99/18.01.7E 5 S. Sh.
33.06.3S/18.01E 4°5 do
33.05.59/18.01.6E 12°5 do.
33.07.99/18.02.1E 2°5 f. S.
33.07.79/18.02.4E 4 S. Sh.
33.07.4.9/18.02.5E 3°5 do.
33.05.79/18.01E 5 Gr. R
Position Depth Bottom
(Metres)
33.49.59/18.27.5E 12°8 S. Sh. R.
33.48.35/18.24E II S. St.
33.4.7.59/18.24.3E 19°5 S. Sh. St.
33.485/18.24.3E 16 Sh. Gr.
33.52.79/18.29.7E 9 S. St.
33.50.19/18.27.8E V7 Sh. R.
33.50.39/18.28E 15°5 R. Sh.
33.51.25/18.27.3E 23 S.R.
33.52.79/18.26.8E 20°5 R.
33.485/18.21E 16°5 Ss.
34.059/18.21E 11 do.
33.529/18.28E 1 S. Sh.
33.48.6S/18.24.6E 15 Sh. St.
do. do. do.
Suips’ HuLts AND EXPERIMENTAL PLATES SUBMERGED IN TABLE Bay Docks (SH)
Code No.
SH. 69
SH. 71
SH. 74
SH.134
SH.168
SH.204
SH.277
SH.324
SH.366
SH.376
SH.393
SH.400
Date
12.11.46
17.4.46
18.4.46
21.1.47
1.4.47
27-5-47
4-9-47
6.2.48
2.12.48
26.1.49
16.3.49
do.
Remarks.
Norfolk from India and east coast of Africa.
Natal—from India and east coast of Africa.
Windward—local wooden yacht 9 months in water.
Empire Liddell—local ship 1 month in water.
Experimental plate submerged for 120 days.
Experimental plate submerged for 175 days.
Experimental plate submerged for 275 days.
Barge working in Table Bay.
Wooden Teredo trap submerged for ? days.
Experimental plate submerged for 96 days.
Wooden frame submerged for 94 days.
Experimental plate submerged for 7 months 2 days.
THE POLYCHAET FAUNA OF SOUTH AFRICA 267
Suip’s HuLts AND EXPERIMENTAL PLATES SUBMERGED IN TABLE Bay Docks (S.H.)
Code No.
SH.415
SH.427
SH.428
SH.430
Date
14.2.49
15.12.49
29-7-53
28.7.58
Remarks.
Wooden frame submerged for over 1 year.
Experimental barge 14 months in water.
Scraping from submerged caisson.
Leeukop—local wooden trawler. 84 months in water.
West Coast Drepcinc (WCD)
No.
WCD. 3
WCD. 5
WCD. 8
WCD.13
WCD.15,
WCD.19
WCD.21
WCD.23
WCD.26
WCD.28
Date
25.2.59
do.
24.3-59
do.
24-459
29-4-59
30.4.59
1.5.59
do.
2.5.59
Position
34.09.8S/18.16.5E
34.099/18.14.8E
34.09.359/18.17.5E
34.09.45/18.16.5E
33-04.39/17-54-7E
33.05.68/17.54.5E
33-04.59/17.55-5E
33.00.48/17.53.7E
33.06.5S/17.55.4E
33.05.59/17.50.4E
FatsE Bay Drepcinc (FB)
No.
FB.301
FB.302
FB.305
FB.306
FB.307
FB.308
FB.309
FB.310
FB.311
FB.312
FB.313
FB.314
FB.316
FB.317
FB.318
FB.319
FB.320
FB.321
FB.322
FB.323
FB.324
FB.325
FB.326
FB.327
‘FB.328
FB.329
FB.330
FB.331
FB.332
FB.333
FB.334
Date
8.7.46
8.9.46
12.11.46
24.11.46
22.2.47
do.
do.
21.447
28.4.47
Position
34.085/18.27E
34.08.59/18.26.5E
34.08S/18.27E
34.09.39/18.27.7E
34.07.59/18.31E
34.08S/18.31.5E
34.07.59/18.29.3E
34.085/18.32E
34.109/18.27.8E
34.09.59/18.27E
34.085/18.29E
34.098/18.27.7E
34.099/18.28E
34.09.59/18.28.3E
34.10.25/18.27E
34.09.25/18.26.8E
34.085/18.29.6E
34.085/18.31E
34.079/18.29E
34.109/18.29.5E
34.099/18.29.5E
34.08.59/18.27E
34.08.99/18.27.4E
34.09.6S/18.26.6E
34.09.85/18.26.1E
34..10.25/18.26.2E
34.10.19/18.26.1E
34.109/18.26.1E
34.09.39/18.26.4E
Off St. James
34.07.59/18.29E
Depth
(Metres)
78
Bottom
Bottom
NNNN
nF nn 49
Tm Fs i
ee os pe
> 9
ee)
a
S. lithoth.
do.
(e)
ANBANNNA
Eee Cx f
a n
268
FAtsE Bay Drepcinc (FAL)
No.
Date
22.2.52
do.
Windmill B
Position
34.09.59/18.35E
34.08.35/18.35.3E
S.E. Oatland Pt.
do.
do.
34.135/18.28E (plankton)
do.
do.
34.139/18.29E
do.
34.139/18.29E
34.125/18.29E
do
34.059/18.44E
34.09.6S/18.49.2E
do.
34.09.39/18.49.6E
do.
34.09.49/18.50.8E
do.
34.09.45/18.50.4E
34.17.59/18.49.2E
34.17.39/18.48.7E
34.17.25/18.49.4E
do.
34.16.59/18.49.5E
do.
do.
34.10.65/18.4.7.3E
34.08.35/18.35.3E
34.09.49/18.50.4E
34.135/18.29E
34.10.65/18.47.3E
34.09.49/18.51.7E
34.09.39/18.51E
do.
do.
Simons Bay
Glencairn (diving)
do.
do.
do.
Oatland Pt. (diving)
do.
do.
do.
Gordons Bay Quay (diving)
Oatland Pt. (diving)
do.
do.
do.
do.
do.
do.
each (diving)
ANNALS OF THE SOUTH AFRICAN MUSEUM
Depth Bottom
(Metres)
35 S. Sh
24. R.
8-9 R.S
do. do.
do. do.
11-12 S.Sh
do. do.
do do.
15-21 §.Sh.R
do. do.
28 S. Sh
33-36 Sh
do. do.
7 S.R
Q21°5 do.
do. do.
18 R.S
do do.
8 R.
do. do.
12 S.R
22 S.
37-38 S.Sh
16-19 iR.
do. do.
14-17. do
do do.
do do.
36 S. Gr. R
24. R.
12 S.R.
15-21 S.Sh.R.
36 S. Gr. R.
1+5-2°5 5S.
8-12 R.S
45 R.
do. do.
do. do.
23°5 Lithoth
oF R.
2-4 do.
2-7 do.
do do.
I-2 do.
0-2 do
do. do.
do. do.
o-4 do
0-5 do
fe) do
4°5-5°5 S.R
Oo. oO
0-3 R
do. do
do. do
FatsE Bay Drepcinc (FAL)
Date
17.11.53
do.
21.11.53
Position
Oatland Pt. (diving)
34.22.18/18.35.2E
do.
34.12.85/18.36.5E
34.07.19/18.35.6E
34.17.6S/18.39.2E
do.
34.09.99/18.42.4E
34.06.8S/18.40.3E
34.07.15/18.35.6E
34.12.45/18.43.5E
do
34.079/18.32.5E
34.13.99/18.31.6E
do.
34.10.59/18.32.4E
34.20.39/18.31.8E
do.
34.17.49/18.31.4E
34.15.39/18.44.8E
34.18.25/18.45.8E
34.21.19/18.46.8E
34.22.79/18.43.1E
34.20.65/18.39.4E
34.23.79/18.40.9E
34.18.59/18.34.2E
do
34.22.59/18.37.3E
Oatland Pt. (diving)
do.
34.079/18.32.5E
34.07.15/18.35.6E
34.079/18.32.5E
34.22.79/18.43.1E
34.23.79/18.40E
Oatland Pt. (diving)
do.
do.
Noah’s Ark (diving)
do
Windmill Beach (diving)
Roman Rock (diving)
do.
34.239/18.36E
(S.A. Museum material)
Depth Bottom
(Metres)
0-2 S.R.
do. do.
do. do.
24 do.
4-6°5 do.
do. do.
4 do.
25 do.
73 S. Sh. Gr
do. do.
46 S. Sh.
(larva trap, night)
62 S. Sh.
do. do.
36°5 Soak
29 S. Sh.
oy S. Sh. R
42 S2 ik:
do. do.
18 S. Sh. R
40 co. S. Sh
do. do.
36 S.
64 do.
do. do.
49 S. R.
48 S. Sh. Gr. R
55 R.
64 do.
79 gn. M.
82 S. gn. M
88 gn. M.
64 S. M. Sh
do. do.
80 S. Sh.
0-3 R.
4 do.
18 S. Sh. R
Ba Tas, do.
18 do.
79 gn. M
88 do.
4575 RK:
2°5-5°5 do.
10°5 S. Sh.
14 S. Sh. M.
11-14 R.
4-5 do.
14-17 do.
do. do.
12-14 do.
do. S. Sh.
do. do.
35 R.
THE POLYCHAET FAUNA OF SOUTH AFRICA
270 ANNALS OF THE SOUTH AFRICAN MUSEUM
Farsz Bay Drepcine (FAL)
No. Date Position Depth Bottom
(Metres)
FAL.303 9.10.02 34.265/18.37E 73 R.
(S.A. Museum material)
FAL.304 15.10.1897 34.09.459/18.49.5E 18 do.
(S.A. Museum material)
FAL.306 11.9.53 34.22.15/18.35.2E 73 S. Sh. Gr
FAL.314 19.4.55 34.09.65/18.27.4E 26 Ss.
FAL.324 6.10.1898 34.185/18. ?E ? ?
(S.A. Museum material)
FAL.327 10.9.57 Kalk Bay 3-4 R.
FAL.328 31.1.59 34.199/18.34.6E 40 S. Sh.
FAL.334 do. 34.159/18.36E 51 co. S. Sh
FAL.338 do. 34.139/18.35E 44 Ss.
FAL.341 do. 34.118/18.35.5E do. f. br. S
FAL.345 do. 34.11S8/18.33.5E 38 f. S.
FAL.347 do. 34.10.85/18.31E 35 do.
FAL.349 do. 34.08.75/18.31.6E 27 w. 9.
FAL.352 1.2.59 34.23.39/18.40.3E 88 gn. M.
FAL.355 24.2.59 34..23.39/18.39.4E 97 S.R.
FAL.357 do. 34.18.85/18.39E 73 co. S. Sh
FAL.359 do. 34.16.85/18.40.9E 62 S. Sh.
FAL.365 252.59 34.09.25/18.46.6E 30 R.
FAL.367 do. 34.11.15/18.46.9E 37 S.R.
FAL.371 do. 34.12.65/18.46.7E 40 R.
FAL.373 do. 34.15.19/18.44.8E 54 S. Sh.
FAL.375 do. 34.16.88/18.42.8E 60 gn. S. Sh
FAL.376 do. 34.18.75/18.37.2E 72 do.
FAL.378 do. do. do. do
MATERIAL From COMMERCIAL TRAWLERS (TRA)
No. Date Position Depth Bottom
(Metres)
TRA. 20 5.5.46 33.48S/17.35E 311 gn. M.
TRA. 21 4.9.46 34.259/18.10E 301 M. R.
TRA. 25 8.4.48 34.305/20.54E 66 S. M.
TRA. 27 21.7.48 34.48S/20.20E 67 do.
TRA. 30 9.11.47 34.499/20.21E 86 M. R.
TRA. 33 20.7.49 34.559/21.10E go S. R.
TRA. 36 21.1.50 34.359/20.50E 73 M. St.
TRA. 40 —.7.50 34.309/20.57E do. do.
TRA. 41 26.7.51 34.315/20.50E 66 S. M.
TRA. 43 ? 29.499/31.48E 770 M.
TRA. 46 24.9.52 31.259/16.20E 366 gn. M.
TRA. 48 do. 33.159/16.00E 415 M. S.
TRA. 52 do. 32.125/16.38E 394 M.
TRA. 54 28.11.52 34.409/21.35E 73 S. R.
TRA. 55 do. do. do. do.
TRA. 56 do. do. do. do.
TRA. 58 26.11.52 34.285/21.45E 70 S. St.
TRA. 62 = 25.11.52 34.309/21.15E 62 S. M.
TRA. 63 3928.11.52 34.268/21.50E 64 S. M. R.
TRA. 68 6.2.53 32.245/18.07E 69 gn. M.
TRA. 69 do. (plankton) 32.455/18.00E 15 S. R.
TRA. 70 do. 32.295/18.02E 27 M.
eA a7 52.53 32.059/18.14E 66 R. S.
THE POLYCHAET FAUNA OF SOUTH AFRICA 271
MATERIAL From COMMERCIAL TRAWLERS (TRA)
No. . Date Position Depth Bottom
(Metres)
TRA. 73 3.2.53 32.068/16.37E 311 gn. M
TRA. 74 52-53 32.059/17.52E 123 do.
TRA. 75 do. do. do. do.
TRA. 77 6.2.53 32.419/18.03E 27 S. M.
TRA. 80 4.2.53 32.239/17.48E 143 gn. M.
TRA. 84 = 13.11.51 32.379/18.17E 6 Ss.
TRA. 85 22.3.53 32.445/18.02E 18 do.
TRA. 86 233.53 32.485/17.58E 9 do.
TRA. 88 do. 32.445/18.00E II do.
TRA. 89 do. 32.459/18.03E 9 S.R.
TRA. 91 15.7.53 33-519/25.50E 46 M.
TRA. 93 —.1.54 35.035/21.50E 110 S.R.
TRA. do. do. do. do.
TRA.102 —.3.56 34.259/21.30E 55 S. R. Polyz
TRA.104 6.8.56 34.31S/19.21E 22 Ss.
TRA.106 do. 34.339/19.19E a7 do
TRA.107 7.8.56 34.105/18.48E 29 f
(surface plankton)
TRA.108 6.8.56 34.339/19.19E 37 Ss.
TRA. 110 8.9.56 34.199/18.32E 58 S. Sh. R
TRA.112 do. 34.199/18.33E 60 S. R.
TRA.113 do. 34.199/18.32E 58 do.
TRA.114 do. 34.199/18.33E 62 do.
TRA.I15 = 29.11.56 34.159/18.43E 54 Ss.
TRA.116 do. 34.115/18.39E 44 do.
TRA. 121 25.1.57 34.125/18.44E 37 S. R.
TRA.122 do. 34.13.59/18.45E 44 S.
TRA.123 do. 34.125/18.45E 40 S. R.
TRA.127 22.2.7 34.199/18.30E 51 do.
TRA.132 —.2.57 34.205/18.30E 55 Phyllochaetopterus tubes
S. Sh. R.
TRA.133 do. do. do. do.
TRA.135 23.2.57 34.195/18.30E 52 do.
TRA.143 27.3.57 34.189/18.31E 51 do.
TRA.151 6.3.58 34.51S/19.55E 22 R.
TRA.152 do. do. do. do.
Drepoinc By S.A. FisHERIES RESEARCH VESSEL Africana II (AFR)
No. Date Positton Depth Bottom
(Metres)
AFR.689 ? 32.36.68/16.44E 391 gn. M.
AFR.691 8.5.4.7 32.385/16.52E 347 Cl. S.
AFR.707 26.5.47 31.409/16.55E 287 d. gn. M
AFR.718 19.6.47 32.099/18.06E 108 do.
AFR.723-5-7 10.8.47 31.309/17.00E 366 ?
AFR.728 15.8.47 31.149/16.36E 272 Polyz. R
AFR.730 do. 31.30S/16.03E 459 y. Cl S.R
AFR.736 17.8.47 30.429/15.59E 201 co. gn. S. Sh
AFR.761 10.9.4.7 30.139/15.18E 260 Gr. S. R.
AFR.773 14.9.47 28.525/14.50E 194 Cl. S.
AFR.775 15.09.47 29.16S/14.48E 238 Cl. S.R
AFR.783 24.9.4.7 32.439/17.31E 222 S. M.
AFR.789 28.9.47 33.059/17.27E 408 bl. S.R
272
ANNALS OF THE SOUTH AFRICAN MUSEUM
Drepcinc BY S.A. FIsHERIES RESEARCH VESSEL Africana II (AFR)
No.
AFR.790
AFR.791
AFR.801
AFR.830
AFR.831
AFR.835
AFR.842
AFR.882
AFR.945
AFR.9g50
AFR.957
AFR.967
AFR.994
AFR.995
AFR.1224
AFR.1335
AFR.1529
AFR.1532
AFR.1535
AFR.1544
AFR.1545
AFR.1554
AFR.1576
AFR.1578
AFR.1579
AFR.1581
Date
28.9.4.7
4.10.47
7.10.47
19.11.47
do.
20.11.47
25.11.47
10.2.48
19.3.48
20.3.48
22.3.48
23.3.48
19.4.48
do.
15.10.48
13.11.48
4.6.49
do.
9-7-49
23-7-49
do.
28.7.49
9-9-49
do.
do.
10.9.49
Position
33.125/17.40E
32.419/17.18E
32.348/17.52E
32.125/18.42E
35-159/18.39E
?35.099/19.02E
34.359/19.18E
34.399/18.42E
36.255/21.08E
36.44S/21.18E
35.139/21.19E
35-079/20.49E
34.359/21.26E
34.299/21.26E
26.345/15.04E
25.519/14.51E
32.409/17.43E
33.125/17.58E
29.099/16.45E
29.179/16.42E
29.099/16.37E
32.059/18.17E
32.285/18.06E
32.305/17.49E
32.259/17.42E
32.225/17.59E
Mosse, Bay Drepcinc (MB)
No.
Date
Position
34.099/22.07.1E
34.04.25/22.13.8E
do.
34.119/22.10.1E
34.08.59/22.07.2E
34.08.85/22.07.3E
34.11.19/22.09.9E
34.08.35/22.09.4E
34.09.39/22.10E
34.10.19/22.07.8E
34.10.19/22.08E
do.
34.08.59/22.08.8E
34.11.39/22.10E
34.115/22.09.gE
34.10.75/22.09.6E
do.
34.04.39/22.13.5E
34.04.15/22.13.9E
34.04.35/22.14.2E
34.04.85/22.13.1E
do
34.09.15/22.07.3E
Depth
(Metres)
Io
19
do.
3
<
Ske ~ PP
Za.
[e)
Q.
aa
N
ks S
me
nm:
mn nn!
—N
TN
a
re)
BRB mw
ae
ye)
THE POLYCHAET FAUNA OF SOUTH AFRICA
Mosset Bay Drepcinc (MB)
Date Position
19.1.56 34.08.6S/22.07.3E
do. 34.09.1S/22.07.2E
do. 34.08.75/22.07.4E
20.1.56 34.11.35/22.06.3E
do. do.
do. 34.059/22.11.8E
do. 34.06.25/22.10.9E
21.1.56 34.11.48/22.10.1E
17.1.56 34.11.35/22.10E
do. 34.11S/22.09.9E
18.1.56 34.04.85/22.13.1E
ALGoA Bay Drepcinc (LIZ)
No.
No.
Date Position
54-54 33-55-79/25-37-2E
do. do.
5-4-54 33.56.1S/25.40E
6.4.54 33.58.1S/25.38.9E
do. do.
do. 33.58.25/25.38.8E
7-4-54 33-58.45/25.40.5E
do. 33.58.59/25.42E
8.4.54 33-585/25.43E
11.4.54 34.00.45/25.44.5E
do. do.
do. 34.00.85/25.42.4E
do. do.
Drepcinc (SCD)
Date Position
15.4.58 35.279/20.10E
18.4.58 34.205/24.40E
do. do.
19.4.58 34.159/25.05E
20.4.58 32.525/28.12.5E
26.5.58 34.07.39/23.23.8E
do. 234.465/23.27E
24.5.58 34.02.59/25.46.5E
23.5.58 33.4.79/26.04E
22.5.58 33.38.65/26.54.7E
21.5.58 33.039/27.50.2E
19.5.58 32.15.29/28.57.7E
do. 31.38.85/29.34.4E
20.8.58 34.018/25.45.5E
19.8.58 33.379/26.56.6E
16.8.58 33.025/27.56.2E
14.8.58 31.57.29/29.36E
5-7-59 33-315/27.14.5E
15.7-59 31.4.1.79/29.33.5E
Depth
(Metres)
Depth
(Metres)
Bottom
pp
a
TN
FERRED
fe)
(e)
nn
ep)
=
I
Bottom
Bottom
BY provers
ge ele
nin ne D
NM
5
glutinous br. M.
grass.
273
and
274 ANNALS OF THE SOUTH AFRICAN MUSEUM
SoutH Coast Drepcinc (SCD)
No. Date Position Depth Bottom
(Metres)
SCD. 74 16.7.59 32.335/28.38E 55 S. M.
SCD. 78 do. 32.379/28.31E 49 br. S.
SCD. 80 do. 32.439/28.28E 58 St. Sh.
SCD. 82 17.7.59 27.549/33.03E 51 br. S. Sh.
SCD. 89 do. 33.039/27.55E 27 R.
SCD. 94, 96 20.7.59 34.215/25.41E 110 Sh.
SCD. 99 21.7.59 34.339/24.01E 130 R.
SCD.100 do. do. do. do.
SCD.103 22.7.59 35.079/22.15E 120 mS,
SCD.105 23.7.59 34.339/21.28E 67 co. S. br. Sh.
SCD.106 do. 34..359/21.10E 67 S. M.
SCD.109 do. 34.359/21.11E 75 co. S. Sh. St.
Family APHRODITIDAE
Subfamily HERMIONINAE
Aphrodita alta Kinberg 1855
Aphrodite alta. Kinberg 1857, p. 2, pl. 1, fig. 1 a-g. Monro 1930, p. 36, fig. 5 a-l.
Aphrodita A near alta McIntosh 1925, p. 18.
Records: AFR.728(1), 835(1).
Notes: In this species the dorsal setae do not project through the felt, the
eyes lack pigment, the ceratophore of the median antenna is short and stout
but bears a very fine ceratostyle which is three-quarters the length of the prosto-
mium. The ventral setae have curved, bearded tips.
Monro (1930) states that the median antenna is short and stout but this
does not agree with Kinberg’s pl. 1, fig. 15, and it is probable that Monro
was describing the ceratophore from which the ceratostyle had fallen. Again
Monro’s fig. 5a shows the stout dorsal setae as quite smooth. In mine they are
covered with minute hairs.
McIntosh’s specimens in the British Museum have been checked as
identical with my own.
Subfamily PoLyNOINAE
Harmothoe aequiseta (Kinberg) 1855
?Lagisca extenuata Ehlers 1913, p. 446.
Harmothoe aequiseta. Augener 1918, p. 137. Day 1953, p. 400.
Harmothoe crosetensis (non McIntosh) Monro 1930, p. 57 (partim).
Records: SB.118(1); TRA.71(1); FAL.16(1), 30(1), 44(5), 51(p), 56(4),
69(p), 80(p), 223(p); MB. 20(1), 57(1), 77(2); KNY.6(1), 11(1), 21(2).
Notes: The specimens reported by Monro (1930) from Simonstown as
as H. crosetensis are definitely H. aequiseta. McIntosh described H. crosetensis as
having fringed elytra but an examination of his type in the British Museum
THE POLYCHAET FAUNA OF SOUTH AFRICA 275
shows that the surface of the elytron is densely covered with soft papillae right
to the edge but the margins are not fringed. Monro’s specimen of H. crosetensis
from the South Shetland Islands agrees with MclIntosh’s type. ‘The specimen
recorded by Ehlers (1913) as Lagisca extenuata was a juvenile and should
probably be referred to H. aequiseta.
Harmothoe africana Augener 1918
Harmothoe africana Augener 1918, p. 139, pl. 2, figs. 15-19, text fig. 6.
Records: FB.316(1).
Notes: Both H. africana and H. goreénsis (recorded below) are very closely
allied to H. aequiseta if they are not mere varieties of the latter. In H. aequiseta
the larger tubercles on the elytra look like straight dark thorns. In H. africana
the larger tubercles are almost cylindrical and end in 2-4 points but there is
considerable variation among the smaller ones.
Harmothoe goreénsis Augener 1918
Harmothoe goreénsis Augener 1918, p. 142, pl. 2, figs. 4-6; pl. 3, fig. 42, text-fig. 7.
Records: LAM.8(1), 22(3), 31(2), 44(1), 47(1), 57(1); SB.180(1), 183(3),
184(3), 189(1), 207(2); T.B.go1(1), 306(1); WCD.8(1), 13(1); FAL.8(1),
58(P), 113(1), 134(5), 144(5), 149(2), 156(6), 216(1), 238(1), 280(4), 338(1),
341(1), 359(1), 375(1); MB.9(1), 13(2), 16(1), 27(3), 41(7), 49(12), 53(12),
56(2), 67(10), 69(1), 74(2); 77(6), 85(2); LIZ.2(6), g(x), 18(4), 35(4), SCD.
40(1), 54(5), 58(5), 74(1)-
Notes: This is a small species first recorded from shallow waters off Angola
and Senegal. This is the first record from South Africa. It differs from H.
aequiseta in having short, blunt, cylindrical or crown-shaped tubercles on the
elytra instead of long sharp ones.
Harmothoe fraser-thomsont McIntosh 1897
Harmothoe fraser-thomsoni. Fauvel 1923, p. 68, fig. 25 a—-e. Day 1953, p. 400.
Records: SB.129(1), 132(1); LB.456(2); LIZ.6(1); SCD.58(1).
Notes: As stated earlier the South African material is a little different
from typical European forms. The elytra have more numerous and crowded
tumid papillae and the notosetae have long naked tips instead of short tips.
Harmothoe gilchristi n.sp.
(Fig. 1 a)
Records: AFR.835(2); FAL.355(1); SCD.22(1).
Description: The holotype is the single specimen from SCD.22. It is complete
and measures 16 mm. by 3:5 mm. with 38 segments. The body is pale but
there are brown markings on the elytra, antennae and dorsal cirri.
276 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 1. Harmothoe gilchristi: a elytron; 6 details of tubercles and papillae; ¢ posterior view of 8th
parapodium; d head; e, e! notoseta; f, f1 neuroseta.
Harmothoe agulhana: g posterior view of 8th parapodium; h, h1 notoseta; 7, 71 neuroseta;
j head; k elytron; / details of ‘tubercles’.
aL
THE POLYCHAET FAUNA OF SOUTH AFRICA 277
The prostomium (fig. 1d) is almost square, slightly broader than long
with very sharp prostomial peaks. The anterior pair of eyes are large and
placed well back on the sides of the prostomium. The three antennae arise
from short swollen cirrophores and all are brown, densely clad with long papillae
and tapering. The median antenna of the holotype is missing but a specimen
from AFR.835 shows that it is twice the length of the prostomium. The laterals
on account of the very large prostomial peaks are markedly ventral in origin
and each is well tapered and three-quarters the prostomial length.
The body is slightly tapered posteriorly and the last 6 segments lack elytra.
The dorsal cirri are tapered, densely papillose and exceed the length of the
neuropodia but not the neurosetae. Each has two dark bands of pigment. The
elytra (fig. 1a) are large, circular, mottled with purplish-brown and cover the
back except for the last few segments. Each has a very small fringe of unicellular
papillae on the external margin and the surface (fig. 1b) is densely covered
with short cylindrical or bollard-like tubercles plus a few elongate papillae.
There are no large posterior vesicles. .
The notopodium (fig. 1c) is well developed with a radiating tuft of
numerous notosetae (fig. 1e). Each is stout and strongly serrated to the blunt
grooved tip. The neuropodium is well developed with a pointed presetal lobe
containing the aciculum and a truncate postsetal lip. The neurosetae (fig. 1f)
are bidentate and have fairly long blades with about 15 rows of well-developed
spinules and long naked tips (fig. 1f). The terminal tooth is broad with a
curved point and the secondary tooth which lies in line with the shaft is
exceedingly long and slender. Its length is almost twice the width of the shaft
at the origin of the tooth and three-quarters the length of the terminal tooth.
H. gilchristi comes fairly close to H. goreénsis Augener but the prostomial
peaks are better developed, the tubercles on the elytra are larger and most
of them are swollen distally instead of being sculptured. Again, the notosetae
have shorter, grooved tips which are rounded not pointed. The secondary
tooth of the neuroseta is also distinctly longer. Type locality: Agulhas bank.
Harmothoe agulhana n.sp.
(Fig. 1 g—l)
Records: ?AFR.7070(1); FAL.365(1); MB.67(1); LIZ.25(1).
Description: The holotype is the single complete specimen LIZ.25 dredged
in Algoa Bay. It is 12 mm. long by 2 mm. excluding setae and has 36 setigers.
The body is narrowly oblong, hardly tapered posteriorly and is pale in alcohol
with a faint network of brown on the exposed parts of the elytra.
The prostomium (fig. 1j) is about as broad as long with poorly marked
frontal peaks. The eyes are rather small and the anterior pair is laterally
situated almost half-way back. The tapered median antenna is as long as the
prostomium but the laterals are very short and stumpy, barely a quarter of
278 ANNALS OF THE SOUTH AFRICAN MUSEUM
the prostomial length. All antennae and cirri are sparsely beset with small
papillae.
' The dorsal cirri are tapered and on all except the last few segments tee
are shorter than the neurosetae. The ventral cirri are very small and distinctly
tapered. The 15 pairs of elytra cover the body except for the last two segments.
Individual elytra (fig. 1k) are large and oval and so thin that the edges tend
to crumple. The colour is generally pale but there is a faint speckling or network
of brown pigment over most of the surface. There is a patch of small rounded
chitinous tubercles on the antero-medial margin, and a scattering of similar
tubercles (fig. 1/) over the surface which do not have chitin-thickened walls
and for this reason are not very obvious.
The notopodium (fig. 1g) is normally developed and the notosetae (fig. 1)
are fairly numerous but rather short and stout and strongly serrated to their
blunt tips (fig. 1h!). The neuropodium is a truncate cone with a small presetal
projection covering the end of the aciculum. The neurosetae (fig. 12) have
blades of normal length with about 10 rows of spinules. Apart from 2-3
superior neurosetae which are unidentate, the tips (fig. 17!) are bidentate
with a strong hooked terminal tooth and a fine secondary tooth.
A fragment of what may be the same species was obtained from station
AFR.707. While generally similar, there are signs of prostomial peaks, the
notosetae are very stout and have grooved tips and the neurosetae have more
rows of spinules, so that both types of setae are very similar to those of H.
gilchristi. But the short lateral antennae and the elytra are very like those of
H. agulhana with only tiny hemispherical tubercles and entire margins.
This species appears to be related to H. ljungmam. The notosetae are
similar and the elytra are not very different though the tubercles are not so
well developed. The neurosetae, however, are quite distinct.
Harmothoe corralophila n. sp.
(Fig. 2 a)
Records : AFR.g50(1) ; WCD.3(1), 13(3); FAL.378(1) ; SCD.100(1), 103(1).
Description: The type was selected from WCD.13. It is 15 mm. long with
37 segments. It is quite white in alcohol with rather glassy setae. The
prostomium is bilobed and broader than long with obvious frontal peaks and
rather large eyes. The anterior pair are half-way back and much wider apart
than the posterior pair. The median antenna is twice as long as the prostomium
and is mounted on a stout ceratophore from which an obvious ridge extends
back along the dorsal surface of the prostomium for half of its length. The lateral
antennae are tapered, markedly ventral in origin and equal to the prostomial
length. All antennae and cirri are smooth. The palps are fairly large.
_ The elytra cover the whole length of the body. All of them are white and
have entire margins without a sign of a fringe but the surface of the first few
THE POLYCHAET FAUNA OF SOUTH AFRICA 279
Fic. 2. Harmothoe corralophila: a first elytron; a’ chitinous tubercle; 5 posterior elytron; 5° soft
papilla; c posterior view of parapodium; d notoseta; ¢ neuroseta; el tip of superior neuroseta;
é* tip of middle neuroseta.
Malmgrenia purpurea: f neuroseta; f 1 tip of neuroseta; g head; h notoseta; h! tip of notoseta;
Jj elytron; k posterior view of parapodium.
280 ANNALS OF THE SOUTH AFRICAN MUSEUM
differs from more posterior pairs which at first sight appear to be quite smooth.
The first pair (fig. 2a) are smaller than the rest, circular in outline and most
of the surface is thickly chitinized and densely beset with numerous highly
chitinized conical tubercles (fig. 2a') plus a few indistinct soft cylindrical
papillae. On some specimens there is only a narrow marginal belt around the
elytron which is not thickly chitinized and free from tubercles but in others
there is a relatively broad naked margin. The second pair of elytra has a more
restricted area of thick chitin and tubercles and a larger area which is naked
apart from the soft digitiform papillae. In the third and fourth pairs there are
even smaller chitinized and tuberculate patches. All succeeding elytra (fig. 2b)
are thin, entirely devoid of tubercles and have only a few indistinct panies
(fig. 251) on the otherwise smooth surface.
The dorsal cirri are smooth and tapered. The first few barely reach the
ends of the neurosetae but further back they are longer. The notopodia are
rather small and the notosetae (fig. 2d) are not very numerous but each is
stout and well developed with widely spaced rows of large serrations or cusps
preceding the long naked tip. Most of the notosetae are sharply pointed but
some of the longer ones have grooved tips.
The neuropodium (fig. 2c) has a pointed presetal lip and a fan of long,
clear, rather slender setae (fig. 2e). The spinules are poorly marked. Although
there are actually 20 rows of spinules which is more than usual, the inferior
neurosetae may at first appear to be smooth because the spinules lie so close
along the blade. The terminal tooth is always well developed but the secondary
tooth is very variable. In superior neurosetae (fig. 2¢!) it is shorter but almost
as broad as the terminal one so that the tip of the blade appears to be bifid.
In middle neurosetae (fig. 2e%) the base of the secondary tooth is stout but the
point is exceedingly fine and often bent or broken leaving a stout stump.
No truly unidentate setae have been seen.
The first specimen obtained (AFR.g50) has lost the anterior elytra and
was provisionally identified as Harmothoe joubint Fauvel 1914 which it resembles
in many respects apart from the anterior elytra and the tips of the neurosetae.
Later several stylasterid corals (Allopora bithalamus) were obtained from
SCD.100 with galls in the shape of open tunnels on their sides. H. corralophila
was found in these galls and is obviously the cause of their formation. Since
both ends of the tunnel are open it is probable that the worm can move in or
out at will and uses the tunnel for protection.
Harmothoe lunulata (Delle Chiaje) 1841
Harmothoe lunulata. Fauvel 1923, p. 70, fig. 26.
Records: FAL.285(1).
Notes: This is a new record for South Africa.
THE POLYCHAET FAUNA OF SOUTH AFRICA 281
Harmothoe saldanha Day 1953
Harnothoe saldanha Day 1953, p. 401, fig. 1 a—d.
Records: SB.202(1); SCD.32(1).
Harmothoe (Lagisca) waahli (Kinberg) 1855
Harmothoe waahli Monro 1933, p. 489, figs. 1-3.
Records: SB.121(4), 127(2); LB.155(2), 161(4); WCD.19(1); FB.302(1),
g11(1), 319(1); FAL.43(2), 51(1), 58(1), 80(1), 235(1), 249(1), 269(1), 367(1);
MB.16(1); LIZ.6(3).
Notes: Several authors have recorded this species under the generic name
Harmothoe but it should be noted that the last 12 segments are narrowed and
not covered by elytra. However I do not feel that this character merits full
generic distinction.
Scalisetosus pellucidus (Ehlers) 1864
Scalisetosus pellucidus. Fauvel 1923, p. 74, fig. 27 a-—f.
Records: LAM.1(1), 8(3), 25(1), 31(2), 63(1); SB.118(1), 119(1), 179(1)5
LB.161(1); TRA.71(1), 102(1); FB.302(1); FAL.8(p), 30(1), 95(P)> 113(p),
122(1), 134(4), 166(1), 184(1), 275(1), 327(1), 338(1), 367(2); MB.4o(1),
74(1), 86(1); KNY.6(1), go(1); LIZ.g(1), 29(1), 35(1); SCD.9(2), 22(2),
100(1).
Antinoe lactea Day 1953
Antinoe lactea Day 1953, p. 403, fig. 2 a-g.
Records: 1.B.299(3), 300(1), 364(5).
Notes: It is surprising that this species has never been recorded outside
Langebaan Lagoon.
Malmgrenia purpurea n. sp.
(Fig. 2 f-k)
Records: WCD.5(1); FAL.229(1), 359(1), 375(1).-
Diagnosis: A purple species with very stout antennae and cirri.
Description: The type material consists of two complete but broken
specimens dredged with Spatangus capensis from stations FAL.359 and FAL.375.
The larger specimen measures 17 mm. and has 38 segments; the smaller
specimen measures 10 mm. and has 37 segments. Both are purple in alcohol
but many of the dorsal cirri and elytra are missing and some of the antennae
as well. For this reason the description is based on both specimens.
The prostomium (fig. 2g) is rectangular and longer than broad with rather
small eyes which are not easily distinguished against the purple background.
282 ANNALS OF THE SOUTH AFRICAN MUSEUM
The anterior pair are set half-way back on the sides of the prostomium. At
first sight it was thought that the insertion of the antennae was harmothoid
and then it was noticed that prostomial peaks are absent and the lateral
antennae are not ventral but subterminal in origin. When the worm was
turned over it was further noted that there is a well developed facial tubercle
and that the bases of the cirrophores which bear the lateral antennae are
fused to the lower side of the prostomium but their distal ends incline upward
so that the antennae appear to be almost terminal. This type of insertion is
best termed.subterminal but it should be noted that it is quite distinct from the
so-called subterminal insertion of Halosydna where the lateral antennae arise
from a lower level than the median antenna only because the latter is actually
dorsal in origin. The median antenna here is terminal and it isa large dark
almost club-shaped organ about as long as the prostomium. Its surface is quite
smooth. The lateral antennae are similar in shape but only half as long. The
palps are rather short and the tentacular cirri are rather stout. The tentacular
segment bears a single seta.
There are 15 pairs of elytra which cover the dorsum. Each is broadly oval
in shape (fig. 27) and quite smooth except for a small patch of minute rounded
tubercles on the anterior margin. There is no trace of a marginal fringe. The
anterior half of each elytron is colourless where it is overlapped by the preceding
one but the exposed posterior half is dark purple with clear cells here and there.
The dorsal cirri (fig. 2k) are similar to the antennae, being dark in colour,
quite smooth and swollen distally before the tip. The notopodium bears about
a dozen stout notosetae and the neropodium which has a pointed or triangular
presetal lip and a shorter, more rounded post-setal lip bears some 20-30
rather short neurosetae. The ventral cirrus is smooth, evenly tapered and
extends almost to the end of the neuropodium.
The notosetae (fig. 2) are stout, very lightly serrated and end in abruptly
pointed tips. Under high power the tiny close-set serrations produce a herring-
bone pattern on the surface of the seta but individual serrations cannot be seen.
The neurosetae (fig. 2f ) have more slender shafts than the notosetae but the
blades are of normal length and bear about 25 rows of very fine, transparent
spinules. The tips (fig. 2f1) are short, the terminal tooth is sharp and well
hooked but the secondary tooth is minute or even absent on some of the inferior
neurosetae.
The pygidium bears a pair of dark sausage-like anal cirri.
Reference to Fauvel (1923) and Monro (1936) shows that the distinction
between the genus Malmgrenia McIntosh 1876 and Eulagisca McIntosh 1885
is not clear. Malmgrenia as defined by Fauvel (1923) is generally similar to
Harmothoe but differs in having the lateral antennae subterminal in origin,
the notosetae very stout and faintly spinulose and the neurosetae either
unidentate or with a minute secondary tooth. Fauvel in his definition states
that the insertion of the antennae is similar to that of Halosydna but his figure
of the head of Malmgrenia castanea shows that it is not similar to that of Halosydna
THE POLYCHAET FAUNA OF SOUTH AFRICA 283
but the same as M. purpurea described above. Eulagisca was not defined by
McIntosh (1885) but has been defined by Monro (1936) in terms which suggest
that it is synonymous with Malmgren. However the figures of Eulagisca
corrientis (the type species of Eulagisca) given by Monro (1930) show that the
insertion of the antennae is similar to that of Halosydna, the notosetae strongly
serrated and the neurosetae unidentate.
The present species M. purpurea may be distinguished from M. castanea
by the position of the eyes, the possession of shorter and much stouter antennae
and dorsal cirri and by the fact that the neurosetae are mainly bidentate.
M. castanea is known to be commensal of the echinoid Spatangus purpureus ;
M. purpurea is probably a commensal of Spatangus capensis.
Lepidonotus durbanensis Day 1934
Lepidonotus durbanensis Day 1934, p. 18, fig. 1 a—c. Day 1951, p. 9.
Records: MB.86(1).
Notes: Only a single small specimen was obtained but it is quite distinct
from the common L. clava var. semitecta. This is the most southerly record of
this Natal form.
Lepidonotus clava (Mont.) var. semitecta Stimpson. 1855
Lepidonotus clava var. semitecta. Willey 1904, p. 256, pl. 13, fig. 4. Day 1953, p. 399.
Records; SB.207(1); LB.161(3); TB.302(1), 305(2),. 309(1),. 313(1),
317(1); TRA.122(1); WCD.8(3); False Bay—5o records on rock o—73 metres,
common; MB.9(2), 16(1), 23(1), 40(12), 49(12), 53(15), 56(2), 59(1), 62(6),
67(x), 85(3); KNY.21(1), 22(1); LIZ.2(2), 6(4), 18(c), 27(3); SCD.89(1).
Polynoe erythrotaenia (Schmarda) 1861
Hemilepidia erythrotaenia Willey 1904, p. 258, p. 13, figs. 6, 26.
Records: SB.179(1); TB.324(1).
Polynoe scolopendrina Savigny 1820
Polynoe scolopendrina. Fauvel 1923, p. 80, fig. 30. Day, 1953, p. 406.
Records: LAM.8(1), 13(1), 25(1), 31(4), 59(common), 63(1); LB.299(2);
TRA.135(1); FAL.r1o(1), 122(f), 219(1), 345(1); MB.41(1), 53(1), 56(1),
67(1); LIZ.2(3), 6(1), 29(2); SCD.58(1).
?Polynoe capensis McIntosh 1885
Polynoe capensis McIntosh 1885, p. 114, pl. 4, fig. 4; pl. 15, fig. 1; pl. 19, fig. 4; pl. ga, figs. 4 and 5.
Notes on the type material. No new specimens have come to hand, but the
type specimens dredged from 98 fathoms off the Cape of Good Hope and
284 ANNALS OF THE SOUTH AFRICAN MUSEUM
now in the British Museum (register number 1885 12.1.94) were re-examined.
The type material consists of blackened fragments of two specimens and several
loose elytra. The median antenna is missing and the lateral antennae which
are terminal in origin, are two-thirds the length of the prostomium. The
anterior pair of eyes are slightly larger than the posterior pair and are dorsal
in position. The total number of elytra originally present cannot now be
determined but one posterior fragment shows 8 posterior segments without
elytra scars. The loose elytra are oval, one half brown and one half white.
The pale half has a triangular patch of small chitinous tubercles. The parapodia
have a fair number of notosetae, each of which is weakly spinulose with an
abruptly tapered tip. The neurosetae are also weakly spinulose and the tip at
first appears to be unidentate, but careful examinations show a small, blunt
secondary tooth.
Although the colouration of the elytra is reminiscent of Polynoe erythrotaenia,
the other characters differ and the terminal insertion of the lateral antennae
shows that this species should be removed from the genus Polynoe. Fresh material
is required before its generic position and exact characters can be determined.
Lepidasthenia elegans (Grube )1840
Lepidasthenia affinis Horst 1917, p. 85, pl. 19, fig. 8.
Lepidasthenia elegans. Fauvel 1923, p. 88, fig. 23 a-g.
Records: TRA.133(1); SCD.58(1).
Notes: These two specimens from dredgings off the Cape and another that
I have seen from the shore of Inhaca Island agree very well with Fauvel’s
description apart from the minor points noted below. They have rather fewer
elytra (23 as against 30-36 for the Mediterranean specimens) and individual
elytra are rather larger though they leave the central third of the back bare
except at the anterior end. Each elytron is speckled with dark pigment except
for a white spot which marks the area of attachment. The prostomium, eyes,
antennae and dorsal cirri are identical. As in the Mediterranean form the
notopodia usually lack setae but careful search revealed that a few feet have a
single minute notoseta with poorly marked serrations. The neurosetae are
variable both along the length of the body and within a single fascicle. The
first few feet have 2-3 slender superior setae with long, coarsely spinulose
blades. The remaining setae of an anterior foot are stouter with short spinulose
blades and strongly bidentate ends though the secondary tooth is markedly
smaller than the terminal one. Further back along the body the slender superior
setae are lacking and a giant brown superior seta appears. Moreover the
secondary tooth is reduced and may be completely lacking so that the giant
seta becomes unidentate, with only a few rows of rather worn spinules.
L. elegans has already been recorded from Zanzibar by Potts (1910) but
he makes no mention of the slender superior setae of anterior feet. Horst’s
description of L. affinis from Lombok in the East Indies leaves no doubt that
THE POLYCHAET FAUNA OF SOUTH AFRICA 285
his specimen is conspecific with mine though it had 40 pairs of elytra. He
separates L. affinis from L. elegans as described by Potts on the relative size of
the elytra and minor differences in the disposition of the setae. I feel, however,
that Potts’s description was too brief and that further specimens have shown
these characters are too variable to warrant specific distinction.
Lepidasthenia brunnea n. sp.
(Fig. 3 a-d)
Records: FAL.352(2).
Description: ‘The type material consists of two broken specimens, the
largest anterior fragment measuring 40 mm. by 5 mm. (including parapodia)
and having 48 segments. The body is light brown in alcohol with colourless
parapodia and the large deciduous elytra are half brown and half transparent.
The prostomium (fig. 3a) is bilobed and almost twice as broad as long.
The anterior pairs of eyes are larger and wider apart than the posterior pair.
The three long smooth antennae are very alike. All arise from short cerato-
phores on the anterior margin of the prostomium and taper slowly towards the
final slender tip. The median, which is a little longer than the laterals is 5-6
times the length of the prostomium and roughly equal in length to the width
of the body. The tentacular cirri are both about as long as the lateral antennae
and each arises from a stout base with a projecting aciculum. The palps are
stout and equal the length of the tentacles.
The body is long and flattened, brown dorsally and pale ventrally. The
elytra are large and overlap to cover the back They are inserted as usual on
segments, 2, 4, 5, 7,9... and alternating segments anteriorly but on every
third further back and, since the body is broken, it is not possible to say how
many there were, but as the anterior fragment has 21 elytra scars there must
have been many more on the complete worm.
Each elytron is oval in shape, very thin and quite smooth. The anterior
half which lies under the preceding elytron is colourless but the exposed
posterior half is pale brown. The dorsal cirri arise from short broad cirrophores
and extend outwards well past the tips of the setae. Each one is quite smooth
and colourless, rounded in section and tapers evenly to a slender tip. The first
few are considerably longer than those from the middle of the body.
The notopodia (fig. 3b) are reduced to tiny pointed lobes lying on the
dorsal surfaces of the neuropodia. Each contains an aciculum but there are no
notosetae. The neuropodia are stout fingerlike organs projecting from the sides
of the body and posterior ones are longer than the body is broad. Each has a
long pointed presetal lip, a rather shorter postsetal lip and between these two
issues a fan of long setae. The ventral cirrus is short and between it and the
‘body the ventral surface of the parapodium bears a single row of about 8
elongate papillae.
286 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 3. Lepidasthenia brunnea: a head; 6 posterior view of parapodium; c¢, c) superior neuroseta;
d, d' inferior neuroseta.
Sthenelais papillosa: e posterior view of parapodium; f head; g elytron; / neuroseta.
THE POLYCHAET FAUNA OF SOUTH AFRICA 287
The neurosetae are characterized by the possession of a superior group
of setae (fig. 3c) with long blades feathered to their fine hairlike tips which end
in minute knobs. A few such setae are present in the anterior feet of many
species of the genus Lepidasthenia but in this species there are many of them
in all feet. The inferior setae (fig. 3d) are stouter and have much shorter
feathered blades which end in bidentate tips. The secondary tooth is half the
size of the terminal one but the feathering comes so close to the end that the
secondary tooth may be obscured. |
This species differs from L. elegans in many respects particularly the
colouration, the persistence of fine unidentate setae, the lack of a giant seta
and the possession of papillae on the base of the neuropodia. In L. maculata
Potts, which has papillae on the ventral surface of the parapodium, the fine
superior setae are very few and only present in anterior segments and the
bidentate setae have very few rows of serrations. L. berkeleyae Pettibone (1948)
is close but the antennae are shorter, there are fewer fine neurosetae and there
are no ventral papillae on the neuropodia.
Lepidasthenia sp.
Records: AFR.790(1).
Notes: An anterior fragment of 32 segments measuring 18 mm. was
obtained. It is completely colourless with rather glassy setae, and large, delicate
translucent elytra.
The general shape of the body, the head and the appendages is similar to
L. brunnea but there are two important differences. There are no papillae on
the ventral surface of the neuropodia and there are no bidentate neurosetae,
only fine setae with long blades feathered to their slender hairlike tips.
Since the fragment consists of only 32 segments with 15 pairs of elytra it is
impossible to say how many elytra are present in the entire worm. Further,
the bidentate type of neurosetae might appear in posterior feet. For these
reasons this is not described as a new species.
Euphione elisabethae (McIntosh 1885)
Euphione elisabethae McIntosh 1885, p. 62, pl. 9, fig. 3; pl. 17, fig. 7; pl. 18, fig. 10; pl. 8A,
figs. 3-6.
Records: AFR.691(1), 707(1), 791(1), 1529(1), TRA.21(1).
Hhololepida australis Monro 1930
Hololepida australis Monro 1930, p. 93, fig. 9 a—h.
Records: AFR.707(1).
Notes: The present material has been compared with Monro’s type in
the British Museum and found to be conspecific. The nuchal flap is triangular.
The elytra which are fragile and deciduous bear numerous three-pronged
288 ANNALS OF THE SOUTH AFRICAN MUSEUM
tubercles exactly as described by Monro. They are reminiscent of the tubercles
of Halosydna and indeed the genus Hololepida is closely related to Halosydna.
The notosetae are not smooth as stated by Monro for under high magnification
they show steplike serrations. The superior neurosetae are as shown by Monro
but his intermediate type was not found. The inferior neurosetae are bidentate
and the secondary tooth, though finer than the terminal one, is epeh oe: as long
so that the ends of these setae seem to be split.
This is a new record for South Africa.
Polyeunoa laevis Mcintosh 1885
Enipio rhombigera Ehlers 1908, p. 47, pl. 4, figs. 1-12.
Polynoe agnae McIntosh 1925, p. 21, pl. 2, figs. 3 and 4.
Hemilepidia erythrotaenia (non Schmarda) McIntosh 1925, p. 26, pl. 2, figs. 9 and 1o.
Records: AFR.789(1), 831(1); TRA.48(1); WCD.3(4).
Notes: The present specimens, like those described by Monro (1936),
have elytra which are smooth apart from a triangular patch of minute
hemispherical tubercles near the point of attachment. These papillae are
absent from MclIntosh’s type. The superior neurosetae have markedly stronger
spinules than the inferior ones. Small specimens of 18 and 20 mm. have a
minute but distinct secondary tooth on the neurosetae but large specimens
are usually unidentate though vestiges of the secondary tooth may occasionally
be found among the inferior setae.
The identity of Ehlers’ Enipio rhombigera with P. laevis has long been recog-
nized. A recent examination of the type of Polynoe agnae (also called Eunoa
agnae by McIntosh 1925) which is now in the British Museum (registered
number 1924:7:21-27) shows that this is also P. laevis. The specimen recorded
by McIntosh (1925) as Hemilepidia erythrotaenia is in a very poor condition but
re-examination again shows that it is also P. laevis.
Subfamily SIGALIONINAE
Pholoe minuta Fabricius var. inornata Johnston 1865
Pholoe minuta var. inornata. Fauvel 1923, p. 120, fig. 44 a—h.
?Pholoe minuta Ehlers 1913, p. 450. Augener 1918, p. 118.
Records: SB.189(1); FAL.22(1), 152(1), 314(1); SCD.26(1).
Notes: The median antenna has a stout base and a slender tip, the whole
equalling the length of the prostomium. The eyes are coalescent. The elytra
are rounded to reniform and the margins carry soft papillae which are not
annulated. The papillae on the neuropodia are not obvious and the shaft-heads
of the neurosetae are lightly serrated. Ehlers (1913) recorded P. minuta from
False Bay and Augener (1918) who recorded the variety inornata from South
West Africa has discussed its distribution and affinities, but the distinctions
between the various species and varieties given by Fauvel (1923) are not very
convincing.
THE POLYCHAET FAUNA OF SOUTH AFRICA 289
Sthenelais boa (Johnston) 1833
Sthenelais boa. Fauvel 1923, p. 110, fig. 41 a-l. Day 1953, p. 406.
Records: LB.299(3); TRA.88(1), 133(1).
Sthenelais limicola (Ehlers) 1864.
Sthlenelais limicola. Fauvel 1923, p. 113, fig. 42 a-g.
Records: AFR.736(1); ?TRA.106D(1); FAL.184(3), 206(1), 228(1),
237(1), 238(3), 242(1), 341(1), 352(7), 375(3), 376(4), 378(1); ?>MB.79(1);
SCD.109(1).
Sthenelais papillosa n. sp.
(Fig. 3eJ)
Records: FAL.223(1), 334(1), 341(1).
Diagnosis: A species with specked elytra lacking sumple serrate neurosetae
and having a papillose ventral surface.
Description: The type material consists of two fragmentary specimens
dredged in False Bay. The larger specimen (FAL.223) is an anterior end of
40 segments and the smaller one FAL.334 is in three fragments but possesses
elytra. It is estimated that the larger specimen might have measured 40 mm.
by 3 mm. when complete.
The prostomium (fig. 3 f) is ovoid, a little longer than broad, with a stout
median ceratophore and two pairs of well-marked eyes. The ‘ctenidia’ on the
ceratophore are rather small and the ceratostyle is slightly longer than the
prostomium. The tentacular segment has a flattened presetal lip, a bundle of
simple notosetae, a short dorsal cirriform appendage (which, according to
Fauvel (1923), corresponds to the lateral antenna) a large dorsal cirrus (or ?
postsetal lobe) and a long ventral cirrus arising from the base of the foot.
The body is elongate and the whole of the ventral surface including the
midventral line and the bases of the parapodia is densely covered with small
spherical papillae. Anterior elytra are not known but those from the middle
of the body (fig. 3g) are reniform without any external notch and the margins
bear minute unicellular papillae which are elongate laterally and spherical
posteriorly. The surface of each elytron is speckled with brown and studded
with tiny, transparent, very lightly chitinized and flattened tubercles or
cushion-like papillae.
The notopodium (fig. 3e¢) has about 6 short stylodes and the usual bundle
of long notosetae. The neuropodium has 2-3 short stylodes at the apex of the
acicular lobe and a low presetal lip edged with about 8 elongate papillae.
The ventral margin of the parapodium as mentioned above, bears numerous
spherical papillae and a single short ventral cirrus.
The notosetae are typical. The neurosetae (fig. 3j) lack superior simple
serrate setae and the compound setae are all very similar. The shafts are stout,
290 ANNALS OF THE SOUTH AFRICAN MUSEUM
the triangular shaft-heads are lightly serrate and most of the blades are long
and simple though some of the short inferior ones have 2-3 poorly marked
articulations. 7
S. papillosa has been compared with the types of S. zeylanica Willey and
also S. variabilis, S. orientalis and S. foliosa Potts (1910) all of which have a
papillose ventral surface but in each case other characters differed. S. papillosa
also approaches §. minor in the lack of simple serrate neurosetae but the latter
may be distinguished by the possession of elongate and _pluriarticulate
neurosetae and the lack of papillae on the ventral surface.
Sigalion squamatum Delle Chiaje 1841
Sigalion squamatum. Fauvel 1923, p. 104, fig. 39 m-—o.
Records: FAL.243(1), 357(1).
Notes : wo anterior fragments were obtained belonging to large specimens
probably exceeding 100 mm. Four tiny well-separated eyes are just visible
through the skin of the prostomium. The dorsal cirrus of the first setiger is
4—2 the length of the ventral cirrus but dorsal cirri are absent from the second
and subsequent setigers. A clavate presetal papilla appears at the end of the
notopodium from the 5th foot onwards. Each elytron is rectangular and the
external margin bears simple papillae on its upper surface and bipinnately
branched papillae along its external margin, each of these having 7—10 pairs
of branches. Rudimentary cirriform gills appear on the medial and lateral
margins of the elytrophore of the second or third foot and by the 6th foot the
gills are well developed. Later the medial gill decreases in size but the lateral
one remains large. The anterior face of the notopodium of each foot has a
patch of conical tubercles near its base.
The notosetae are numerous and minutely serrate. The neurosetae are of
5 types: (a) about 6 simple bipectinate setae in the superior group above the
aciculum. (5) About 2 compound setae with coarsely serrate shafts and tapered,
pluriarticulate blades. (c) About 4 compound setae with swollen, closely
serrate shaft-heads and pluriarticulate blades. (d) About 6 compound falcigerous
setae with fine serrations on the shaft-heads and simple bidentate blades.
(e) Very numerous inferior compound setae with very lightly serrate shaft-heads
and long pluriarticulate blades. All the various types of compound setae of
this and all other species of Sigalion which have been examined have bidentate —
tips. Statements to the contrary are probably due to the examination of
broken-tipped setae.
This South African material has been compared with specimens of
S. squamatum from Naples, which is the type locality. The European material
shows that the development of the superior ‘stylode’ (=presetal lobe) of the
neuropodium is variable and not of much value in separating S$. squamatum
from S$. mathildae. Again the pennate marginal papillae on the elytra are not
normally as widely different as figures 39 c and m in Fauvel (1923) would
THE POLYCHAET FAUNA OF SOUTH AFRICA 291
suggest. I agree with Fauvel that the main difference lies in the setae, though
the tubercles on the face of the notopodium are more poorly developed in
S. mathildae. Incidentally these same tubercles are exceedingly long and
occasionally branched in S. buskiz McIntosh (1885) and suggest that this is a
valid species which lacks falcigers with simple bidentate blades. It is also to be
noted that Fauvel makes no reference to the medial gill on the elytrophores,
and this character was not checked on the Naples material.
Sigalion capense n. sp.
(Fig. 4 af)
Records: FAL.237(1), 375(1); MB.4(1).
Description: FAL.237 an ovigerous female, is selected as the type. It is
16 mm. long by 2 mm. wide and is broken at the 6oth segment. It is quite white
in alcohol.
The prostomium (fig. 4a) is almost oblong, a little longer than wide and
somewhat rounded posteriorly. The two pairs of small eyes are visible through
the skin about the middle of the prostomium. The anterior and posterior pair
are close together on either side. The antennae are small cylindrical papillae
arising from the prostomium at its junction with the forwardly projecting
tentacular segment which bears two pairs of subequal tentacular cirri. The
palps are long and slender. The ventral cirrus of the second foot is about the
same length as the tentacular cirri. A single cirriform branchia arises from the
lateral side of the elytrophore of the 4th and all succeeding segments and in
anterior segments there is also a small branchia (or ctnidium) on the medial
side of each elytrophore.
The posterior elytrophores are swollen with developing eggs. The elytra
themselves (fig. 4c) are rounded to rectangular with smooth surfaces and bear
10-12 bipinnate papillae on the external margin; each of these has 4-8 pairs of
branches (fig. 4c).
The notopodium (fig. 4b) is swollen distally with a single large presetal
papilla (stylode) at its end and three glandular cushions on its superior margin.
It bears a bundle of long slender setae with hairlike tips. The stouter ones are
serrate on the inferior margin.
The neuropodium is obliquely truncate with a bluntly conical acicular
lobe, a vestigial presetal lip, a well-developed, trangular postsetal lip and a
long tapered ventral cirrus. There are 4 types of neurosetae. The supra-acicular
neurosetae include: (a) 3-6 simple bipinnate setae (fig. 4d); (b) 4 fairly stout
compound setae (fig. 4) with long pluriarticulate blades and serrated shaft-
heads; (c) about ro fairly stout compound setae with long pluriarticulate
blades and smooth shaft-heads (fig. 4¢) The infra-acicular setae include: (d)
about 4 setae similar to group (c) and (e) very numerous fine compound setae
with long pluriarticulate blades and smooth shaft-heads. It is emphasized that
292 ANNALS OF THE SOUTH AFRICAN MUSEUM
h
Fic. 4. Sigalion capense: a head; b parapodium; ¢, c! elytron and details of marginal papillae;
d simple serrate neuroseta; e smooth-shafted neuroseta; f serrate-shafted neuroseta.
Psammolyce articulata: g head; h neuroseta; j parapodium; k, k! 6th elytron and details of
margin; / posterior elytron.
2
‘
THE POLYCHAET FAUNA OF SOUTH AFRICA 293
all compound setae have long pluriarticulate blades with minutely bidentate
tips’: .
S. capense may be distinguished from S. sqguamatum and S. mathildae by the
lack of neuropodial setae with simple blades. In this and other respects it
resembles S. ovzgerum Monro (1924) but the latter carries eggs in its swollen
elytra and has supra-acicular neurosetae with excavated shaft-heads as figured
by Monro (1936) fig. rad.
Psammolyce articulata n. sp.
(Fig. 4 g-l)
Records: FAL.117(1), 211(2), 214(1).
Description: Four broken specimens were obtained and specimen FAL.117
which is in two fragments totalling 50 mm. by 4 mm. for 100 segments was
selected as the holotype. It is sandy brown in alcohol with darker setae. The
elytra and the uncovered part of the back between them is covered with sand-
grains and foraminiferan shells and the ventral surface is completely covered
with small rounded tubercles.
The head (fig. 4g) is protected between, but not fused to, the forwardly
directed first pair of feet which bear the tentacular cirri. The prostomium
itself is as broad as long and bears two pairs of large eyes and three antennae.
The first pair of eyes are anterior and ventral and concealed beneath the large
ceratophore of the median antenna. The posterior pair of eyes is on the sides
of the prostomium immediately behind the short stumpy lateral antennae which
are quite separate from the tentacular segment. The median antenna is borne
on a large ceratophore which curves down like an elephant’s trunk.
The first setiger or tentacular segment has well-developed dorsal and
ventral rami and two bundles of setae. The notopodium has a rather short
dorsal tentacular cirrus and a stumpy setigerous lobe. The neuropodium has a
presetal bract, a bundle of neurosetae and a large cirriform postsetal lobe below
which is the long tapering ventral tentacular cirrus. The second segment bears
the first pair of elytra, a short cirriform gill, two setigerous lobes and a long
ventral cirrus. There is no dorsal cirrus. The third segment bears a long stout
elytrophore extending almost to the end of the neuropodium and a shorter
cirrus.
The elytra (figs. 4 k-l) vary along the length of the body but all of them
are covered with sand grains. The anterior pair are pear-shaped and extend
forward over the head. The margins appear smooth and there are certainly
no incisions or large projections. The second pair are reniform. The margins
bear minute papillae and the surface is studded with microscopic tubercles.
The next few elytra are more rounded (fig. 4k) straight in front and with two
short lobes on the medial margin and small pear-shaped lappets posteriorly.
The surfaces are covered with microscopic tubercles and the papillae are better
developed, both on the surface and the margins. All of them are clearly jointed.
294 ANNALS OF THE SOUTH AFRICAN MUSEUM
Further back the two antero-medial lobes tend to disappear (fig. 4/) and the
4-5 posterior lappets become irregular. Most of the elytra are roughly triangu-
lar and fairly small so that the middle third of the back is uncovered.
The feet (fig. 43) show little change from the 4th onward. The notopodium
is well developed. The neuropodium is obliquely truncated with a presetal
row of elongate papillae and tufts elsewhere. The lower surface is covered by
the rounded tubercles which extend on to the parapodia from the ventral
surface. The ventral cirrus is long and tapered and 2-3 very long and slender
papillae arise from its base.
The notosetae are numerous and very fine. Each is minutely serrated to
its hairlike tip. The neurosetae (fig. 4h) are all compound and falcigerous.
Those of the second setiger have densely serrated shafts but in later feet the
serrations are reduced to 3—5 rows on the shaft-head. The blades are all
bidentate but never pluriarticulate.
In his diagnosis of the genus Psammolyce, Fauvel (1923) states that the
lateral antennae are fused to the first setiger or possibly absent in P. inelusa.
Monro (1936) makes the same remark. In the present species as stated, the
lateral antennae arise from the prostomium which is quite separate from the
first setiger. P. articulata is also characterized by the possession of jointed papillae
on the elytra. Faint indications of jointing were seen in the papillae of P.
semiglabra Monro (1936), the type of which was examined, but the two species
differ in many characters. Possibly P. articulata comes closest to P. zeylanica
Willey (1905) in the shape of the elytra but the latter again has the lateral
antennae fused to the first setiger.
Thalenessa oculata (Peters) 1854
Euthalenessa dendrolepis (Clap.) Fauvel 1923, p. 114, fig. 42 h-o.
Euthalenessa oculata Day 1953, p. 407.
Records: SB.207(1); TB.301(1), 304(4); FB.312(1), 321(1); FAL.233(1),
238(1), 349(3); SCD.105(2 juveniles).
Family CHRYSOPETALIDAE
Bhawania goodei Webster 1884
Bhawania goodei. Augener 1918, p. 98, pl. 2, figs. 1-2, text-fig. 1. Day 1953, p. 407.
Records: False Bay —21 records on rocky or shelly bottoms between o and
36 metres. MB.13(1), 53(1), 56(1), 77(1), 85(1), 86(1); LIZ.9(1), 27(1),
33(1); SCD.58(1), 89(1).
Paleanotus chrysolepsis Schmarda 1861
Paleanotus chrysolepis Schmarda 1861, p. 163, pl. 37, figs. 326-9. Ehlers 1913, p. 450. Day
1957, p. 66.
Records: False Bay—15 records on rock or shelly bottoms between o and
40 metres.
THE POLYCHAET FAUNA OF SOUTH AFRICA 295
Family AMPHINOMIDAE
Chloeia inermis Quatrefages 1865
Chloeia gilchristi McIntosh 1925, p. 15, pl. 1, figs. 7-8. Day 1934, p. 27, fig. 4 a-b.
Chloeia inermis. Monro 1936, p. 80.
Records: AFR.801(1); TRA.20(1).
Notes: The specimens were recovered from the stomachs of fish and are
rather soft but they still retain rather vague purplish markings along the
middorsal line and the dorsal cirri are purple. Monro states that the gills begin
on setiger 5 but here they begin on setiger 4. The setae agree perfectly with
Monro’s description. A spur is virtually absent from most of the setae though
a minute one can be seen on some of the ventral ones. All but a few of the
harpoon setae are smooth.
Euphrosyne capensis Kinberg 1857
Euphrosyne capensis. McIntosh 1885, p. 1, pl. 2, fig. 5, pl. 1A, figs. 1-3. Day 1953, p. 408.
Records: LAM.8(1), 15(1), 31(8), 35(2), 47(1), 51(3), 57(1), 59(3), ©3(1) ;
TB.305(1), 323(1); False Bay—17 records between o and 42 metres on rock
or broken shell; MB.16(2), 49(2), 67(1), 77(1), 78(2); LIZ.18(1)
Euphrosyne myrtosa Savigny 1818
Euphrosyne myrtosa. Gravier 1901, p. 254, pl. 10, figs. 147-9.
Records: TRA.132(1), 135(1); SCD.40(1).
Notes : Ehlers (1913) who previously recorded this well-known species from
the Cape was doubtful whether it was distinct from E. capensis. Many specimens
of varying size were therefore examined and it appears that there is a constant
difference in the branchiae. In E. myrtosa there are 6-8 branchial trunks and
the tips of the branches are blunt and not expanded. In E. capensis there are
10-11 branchial trunks and the tips of the branches are swollen and pointed
rather like acorns.
Eurythoe chilensis Kinberg 1857
Pareurythoe chilensis Hartman 1948, p. 45, pl. 5, fig. 11.
Eurythoe chilensis Kinberg 1857, p. 13. Monro 1930, p. 28, fig. 1 a—e.
Records: FAL.209(1).
Notes: The single specimen is 20 mm. long. The caruncle is attached to
the dorsum as far back as the second setiger, but a free posterior projection
extends back to the fourth setiger. This species is easily distinguished from the
tropical E. complanata which extends down the Natal coast by its smaller size
and the fact that all the spurred setae are lightly serrated in E. chilensis and
smooth in E. complanata. This is a new record for South Africa.
296 ANNALS OF THE SOUTH AFRICAN MUSEUM
Family PHYLLODOCIDAE
Phyllodoce (Anaitides) madeirensis Langerhans 1879
Phyllodoce (Anaitides) africana Augener 1918, p. 171, pl. 2, fig. 25; pl. 3, figs. 49-51; text-fig. 11
(partim).
Phyllodoce (Anaitides) madeirensis. Fauvel 1923, p. 150, fig. 53 a-d.
Phyllodoce patagonica (non Kinberg) Monro 1930, p. 72 (partim).
Records: AFR.707(1); TRA.56(1), 133(1); FB.316(1); FAL.241(1), |
373(1).
Notes: By the kindness of the Director of the Hamburg Museum I was |
able to examine Augener’s specimens of P. africana from Goree (number
V.1986). There are two specimens, one with the proboscis extruded and one
with the proboscis retracted; both were quite pale in alcohol, the pigmentation
to which Augener refers having faded. The former specimen with the extruded
proboscis is clearly on Anaztides with 6 lumpy ridges on the distal part of the
proboscis and 6 lateral rows each with 8-10 compressed papillae basally. The
prostomium is cordate with 4 normal antennae, a pair of large eyes and an
occipital papilla in the posterior notch. The first tentacular segment is invisible
dorsally but the second and third are distinct. There is no parapodium or
: I I
setae on the third tentacular segment, the formula being 1 + o—_+o0—.
| I N
Anterior dorsal cirri are broadly lanceolate but later ones are obliquely
truncate near the tip and rhomboidal. The ventral cirri are pointed and
longer than the setigerous lobes. This specimen seems to me to be a typical
P. madeirensis.
The second specimen with the retracted proboscis was dissected and its
proboscis proved to be irregularly covered with large pointed papillae except »
at the base where the dorsal wall was bare. The prostomium is elongate and
deeply incised posteriorly forming a pair of lateral lobes which extend back to
segment 2. No occipital papilla was seen. The first tentacular segment is not
visible dorsally, the second is partly visible between the posterior lobes of the
prostomium but the third segment is fully visible. A small setigerous lobe
bearing 2-3 setae is present on tentacular segment 3, the formula thus being
I I ce
1 + o— + S—. Anterior dorsal cirri are broadly lanceolate, almost cordate,
I N
but later ones are longer and more asymmetrical with broad cirrophores.
The ventral cirri are pointed and about as long as the setigerous lobes. ‘This
specimen does not belong to the sub-genus Anaitides and should be named
Phyllodoce (Phyllodoce) africana.
I have also examined the specimen reported by Monro (1930) from
Simonstown under the name of P. patagonica. It is quite clearly P. madeirensis
and may be distinguished from P. patagonica by the absence of setae on tentacular
segment 3.
THE POLYCHAET FAUNA OF SOUTH AFRICA 297
It may also be mentioned that some of my own specimens recorded above
from deeper dredgings are blotched with dark pigment and some of the dorsal
cirri are black and others white. Otherwise they are indistinguishable from the
normal green variety of P. madeirensis.
Phyllodoce sp.
Records: ‘TRA.133(1).
Notes: Specimen TRA.133.L certainly belongs to a species which has not
been recorded from South Africa before but the preservation is not too good
and its exact determination is doubtful in consequence.
The body is brownish blotched with darker pigment. The proboscis
appears to be diffusely papillose. The prostomium is cordate and there is a
small occipital button in the posterior notch. The first tentacular segment is
fused to the prostomium but the second and third are distinct. The tentacular
cirri are unusual for they are stout, sausage-shaped and constricted basally.
: I I
The tentacular formula is 1 + 0— +S—. The dorsal cirri are ovoid and
I I
swollen and the oval ventral cirri are a little larger than the blunt setigerous
lobes. The setae have oval shaft-heads and short blades.
Phyllodoce macrophthalma Schmarda 1861
_ Phyllodoce macrophthalma. Fauvel 1923, p. 146, fig. 51 f-g.
Records: MB.20(t).
Notes: The single 12 mm. specimen is referred to P. macrophthalma with
some hesitation. The body is slender and dark green. The prostomium is
cordate with a very small posterior notch and the presence of an occipital
button is doubtful. The proboscis on dissection seems to be lightly papillose.
The first tentacular segment is fused to the prostomium but the second and
third are distinct. All the tentacular cirri are well developed and cylindrical,
the formula being: 1 + ele = ey
I I
The dorsal cirri are cordate, possibly a little longer than broad and the
ventral cirri are ovoid. The setigerous lobe has a notched presetal lip and
bears numerous setae with oval shaft-heads striated distally and fairly short
blades. |
Schmarda (1861) stated that the dorsal cirri are rhomboidal but both Ehlers
(1913) who recorded this species from Simonstown and Fauvel (1923) follow
Saint-Joseph (1888) and describe cordate dorsal cirri. I have not seen the latter
paper.
298 ANNALS OF THE SOUTH AFRICAN MUSEUM
Phyllodoce (Anaitis) capensis n. sp.
(Fig. 5 a-c)
Records: FAL.316(1); TRA.133(1).
Description: Specimen FAL.316 which measures 35 mm. by 1:5 mm.
for about 100 segments was chosen as the holotype. It is depressed and tapered
posteriorly and is creamy white in alcohol.
The prostomium (fig. 5a) is broadly rounded anteriorly and produced
posteriorly. The frontal antennae are well developed, the eyes are large and
there is a small occipital button. The first and second tentacular segments are
fused and form a sort of transparent shield which grows forwards over the sides
of the prostomium to cover part of the eyes but the occipital button in the
mid-dorsal line is not covered. The third tentacular segment is distinct. All the
tentacular cirri are cylindrical and tapered but T, and V, are shorter than
T, and T,. There is a small setigerous lobe on the third tentacular segment
: I I
so that the tentacular formula is: 1 + o— + S_.
I
The proboscis was not extruded and dissection was not entirely successful.
The distal part definitely has 6 longitudinal rows of large soft papillae or
rugosities but the oral end is indistinct. There were no obvious rows of papillae
and it may be smooth.
The papapodia and dorsal cirri (fig. 55) are essentially similar throughout
the length of the body. The dorsal cirri are fairly large and rounded to broadly
cordate but they do not cover the back. The ventral cirrus is oval and larger
than the setigerous lobe which has a notched presetal lip and bears about 12
fine heterogomph spinigers. The shaft-head (fig. 5c) is asymmetrical with a
large curved tooth accompanied by 3-4 smaller denticles on one side and a
smaller tooth on the other side. The finely serrated blade is long, fairly broad
basally and tapers gently towards the tip.
This South African species is obviously related to P. (A.) kosteriensis
Malmgren from northern Europe but differs in several respects. The shield
formed by the fusion of T, and T, is better developed, the proboscis is different,
the dorsal cirri are proportionately longer and the setae have slightly different
shaft-heads. P. (A.) wahlbergi Malmgren from the Arctic is, according to
Bergstré6m (1914) a much broader worm with a single blunt tooth on the
shafthead of the seta.
Phyllodoce castanea Marenzeller 1879 —
Genetyllis castanea Bergstrom 1914, p. 158, pl. 3, fig. 4, text-fig. 53.
Phyllodoce rubiginosa Augener 1918, p. 168.
Records: SB.120(1); FB.302(2), 307(3); FAL.128(1), 266(1); MB.23(1).
299
THE POLYCHAET FAUNA OF SOUTH AFRICA
—
Fic. 5. Phyllodoce (Anaitis) capensis: a head; 6 anterior view of parapodium; ¢ seta.
Eulalia bilineata: d head; e anterior view of parapodium; / seta.
Eulalia macroceros: g head; h anterior view of parapodium; 2 seta.
300 ANNALS OF THE SOUTH AFRICAN MUSEUM
Notes: The Director of the Hamburg Museum very kindly sent me the
specimen V.8731 recorded by Augener 1918 from Swakopmund under the
name of Ph. rubiginosa. The proboscis was retracted but when dissected proved
to be very long and covered with irregularly arranged conical papillae. The
whole specimen is rusty red all over with an ovoid prostomium not notched
posteriorly and fused to tentacular segment 1. The antennae are very small.
All the tentacles are short, tapered and rounded. Tentacular segment 1 is not
visible dorsally but 2 and 3 are distinct and bear setigerous lobes with numerous
setae the formula being 1 + Gib + s—. Dorsal cirri are broadly cordate and
I
almost symmetrical. Ventral cirri are ovoid to reniform and extend beyond
the rounded setigerous lobes.
P. castanea and P. rubiginosa are very alike but the latter is described by
Fauvel as having large not small antennae and he figures the dorsal cirri as
asymmetrical. Certainly Augener’s specimen from Swakopmund is very like
mine from the western and southern coasts of the Cape.
Eulalia (Steggoa) capensis Schmarda 1861
Eulalia capensis Schmarda 1861, p. 86, pl. 29, fig. 231. Willey 1904, p. 259.
Eulalia viridis var. capensis McIntosh 1904, p. 34. Day 1953, p. 30.
Eulalia viridis (non Muller) Ehlers 1913, p. 455. Day 1934, p. 30.
Steggoa magalhaensi (non Kinberg) Augener 1931, p. 284.
Records: LAM.18(1), 59(3), 63(1); FB.302(1), 326(1); FAL.23(1), 29(1),
44(2); 69(1), 106(1), 149(1), 245(1), 334(1); TRA.123(r).
Notes : This species although generally similar to E. viridis differs in having
a more flattened ventral tentacular cirrus on the second tentacular segment and
in lacking setae on the same segment so that the formula is 1 + 07 “ Si",
I
It thus belongs to Bergstrém’s genus Steggoa here relegated to a sub-genus.
On the other hand I cannot agree with Augener (1931) that it is identical
with Steggoa magalhaest Kinberg which has spear-shaped dorsal cirri three times
as long as broad.
Eulalia (Hypoeulalia) bilineata (Johnston) 1840
(Fig. 5 df)
Records: SB.197(1); FAL.371(1); MB.66(2), 87(2); SCD.40(3).
Description: These South African specimens differ in several important
respects from the descriptions given by Bergstré6m (1914), p. 165, text-fig. 57,
and Fauvel (1923), p. 162, fig. 58 a—e, which themselves differ in minor respects.
For this reason a full description is given below. The largest of the 3 specimens
from SCD.40 measures 20 mm. by 1 mm. for 155 segments. It is a slender
yellowish worm with two dark green stripes along its back just above the
parapodia. These colours persist in alcohol.
THE POLYCHAET FAUNA OF SOUTH AFRICA 301
The prostomium (fig. 5d) is rounded in front and almost straight
posteriorly. The frontal antennae are well developed and the median antenna
arises well in front of the eyes which are relatively large. The proboscis is
covered with small conical papillae. The relation between the prostomium and
the first tentacular segment is difficult to ascertain. In contracted specimens
the two seem to be fused but in expanded specimens the first tentacular segment
seems to be reduced dorsally but not actually fused to the prostomium. The
second and third tentacular segments are definitely distinct. The three dorsal
tentacular cirri are cylindrical and those on the second and third segment are
rather long; on the other hand V, is short and flattened though not bladelike
as in the sub-genus Sige. The second and third tentacular segments have
: I I
setigerous lobes with setae so that the tentacular formula is 1 + S— + S—.
I
The body segments do not change appreciably along the length of the
worm. The dorsal cirrus (fig. 5¢) is bluntly lanceolate in adults but distinctly
broader, almost cordate in juveniles. The setigerous lobe is bluntly rounded and,
as usual, the presetal lip is deeply notched. The ventral cirrus is ovoid. ‘The 12-
16 setae are heterogomph spinigers (fig. 5f) with ovate shaft-heads striated
distally and lightly serrated blades of normal length.
According to Bergstrém, Hypoeulalia bilineata has the first segment fused to
the head, V, is cylindrical and the dorsal cirri are ovoid. According to Fauvel
(1923) Eulalia bilineata has the first segment narrowed but fairly short and the
dorsal cirri are stout and oval-obtuse. Judged by these descriptions the most
‘marked difference between South African and European specimens concern
the shape of V,.
This is a new record for South Africa.
Eulalia (Eumida) sanguinea (Oersted) 1843
Eulalia (Eumida) sanguinea. Fauvel 1923, p. 166, fig. 59 f-k.
Records: LAM.22(1); LB.161(2), 382(1); FB.316(1); FAL.70(3), 376(1);
MB.86(1); SCD.9(3), SCD.109(1).
Eulalia (Pterocirrus) macroceros Grube 1860
(Fig. 5 g-4)
non Sige macroceros Bergstrém 1914, p. 136, text fig. 40.
Eulalia (Pterocirrus) macroceros Fauvel 1923, p. 167, fig. 60 d—g (partim).
Eulalia (Pterocirrus) ?macroceros Day 1953, p. 411.
Records: MB.66(1), 86(1).
Description: The discovery of two further specimens allows me to confirm
my previous identification, and to complete the description.
The body is broad and short and dark green in life but brownish in alcohol.
The prostomium (fig. 5g) is cordate with a large posterior excavation containing
a dark cushionlike lobe which may represent the dorsal remnant of the first
302 ANNALS OF THE SOUTH AFRICAN MUSEUM
tentacular segment. The eyes are large and there are usually pigment granules
behind them. The two pairs of frontal antennae are surprisingly long and the
dorsal antenna arises slightly in front of the eyes. The proboscis has not been
seen everted but on dissection the base proves to be smooth but further along
there are large soft rugosities. It is certainly not densely covered with cylindrical
papillae as in Eulalia viridis or E. capensis.
The first tentacular segment is fused to the prostomium and bears a pair
of cylindrical tentacular cirri. The second segment is distinct and bears a pair
of long cylindrical dorsal tentacular cirri on swollen cirrophores, a pair of
flattened, often blade-like ventral cirri but there is neither setigerous lobe nor
setae. The third tentacular segment bears a long cylindrical dorsal cirrus, a
normal foliaceous ventral cirrus but no setigerous lobe or setae. The tentacular
formula is thus 1 + lia + Ges.
I N
The parapodia (fig. 52) are very similar throughout the length of the body.
The dorsal cirri are elongate-cordate and pointed, the setigerous lobe has the
usual bilobed presetal lip but here the larger superior lobe is pointed and the
ventral is best described as orbicular with a pointed end. The setae are very
numerous (c.40). Each has a very slightly expanded shafthead faintly striate
distally (fig. 52) and a blade of normal length which is broad and almost smooth
basally and then suddenly narrows and becomes strongly serrated.
As will be seen, the above description does not agree with that given by
Bergstr6ém (1914) and differs in several respects from that given by Fauvel
(1923). In a private communication Dr. K. Banse has informed me that my
(1953) description agreed almost exactly with his specimen from Naples, and
went on to say that EF. macroceros from the Mediterranean (type locality
Quarnero in the Adriatic) is not synonymous with the boreal species E. (Sige)
fusigera Malmgren (1865) (described from Koster-Inseln and Skelderviken in
Sweden and Drobnak in Norway).
Bergstrém’s description is based on the Swedish material and should be
referred to E. (S.) fusigera. Fauvel presumably had more than one species
before him.
Some doubt remains regarding the subgenus to which EF. macroceros
should be referred. It does not fit exactly into any of the numerous genera
used by Bergstrém, and certainly cannot be referred to Sige which has setae
on the second and third tentacular segments. It is suggested here that it should
be referred to Claparéde’s Pterocirrus established for P. velifera Claparéde (1865)
from Naples which according to Grube (1880) is synonymous with E. macroceros.
If this be accepted Pterocirrus should be defined as a subgenus of Eulalia with the
first tentacular segment fused to the prostomium which has a posterior
excavation containing a cushion-like lobe. The second and third tentacular
segments are distinct, and tentacular cirrus V, is flattened. There are no
, j ! I
setae on any of the tentacular segments, formula being: 1 + o__ + 0—.
I N
THE POLYCHAET FAUNA OF SOUTH AFRICA 303
Eulalia (Sige) falsa n. sp.
(Fig. 6 a—c)
Records: FAL.187(2), 338(1); MB.87(1).
Description: ‘The type material consists of the two specimens from FAL.187
dredged in False Bay. One is a regenerating individual 17 mm. long by 1-5 mm.
wide with 60 segments and an everted proboscis. It is brownish in alcohol.
The other is a juvenile 9 mm. long by 1 mm. wide with 60 segments. A fresh
specimen (FAL.338) has a brown dorsum with a paler head and para-
podia.
The body is elongate and of the usual proportions. The prostomium
(fig. 6a) is cordate with a pair of large eyes, subulate frontal antennae and a
median antenna which arises between the eyes. The first tentacular segment is
fused to the head but there is an area between the posterior lobes of the
prostomium which may represent the dorsal remains of this segment, or an
anterior projection of segment 2. The second and third tentacular segments
are distinct and bear long, dorsal, tentacular cirri. Tentacular cirrus V, is
flattened and may even be blade-like in juveniles. Both the second and third
tentacular segments bear setae but those on the second segment are very few
and arise from the ventral cirrophore, there being no separate setigerous lobe
on this segment. A small setigerous lobe with numerous setae is present on the
third segment and below this is the foliaceous ventral cirrus, the tentacular
formula being: 1 + ou + S_". The extruded proboscis is faintly hexagonal
I N
with 6 low longitudinal ridges. The surface is covered with very small and poorly
marked papillae so that on first inspection it appears to be smooth. The opening
is encircled by 20 large rounded papillae.
The parapodia of the adult (fig. 65) are very similar throughout. The dorsal
cirrus is elongate-cordate with a pointed apex, and in specimen MB.87 there is a
dark central spot. The setigerous lobe has a bilobed presetal lip, the superior
lobe being large and pointed and the inferior lobe being small and blunt.
In the juvenile specimen these characters are more marked and the superior
lobe has a very long pointed lobe. The setae (fig. 6c) are numerous fine
heterogomph spinigers with very slightly expanded shaft-heads bearing about
4. small denticles at the distal end. The blades are smoothly tapered and finely
serrated.
This species is rather similar to Bergstrém’s description of Sige macroceros
which, as shown above, really refers to E. (S.) fusigera. However there are
differences in the shape of the prostomium, the nature of the proboscis, the
shape of the cirri and the structure of the setae. Later work may show that this
South African material is conspecific with E. (S.) fusigera from northern Europe,
but to avoid further confusion in the synonymy it is as well to keep them
separate at present.
304 ANNALS OF THE SOUTH AFRICAN MUSEUM
ove
ig
tee
RRO E HAIN
Pine ;
Fic. 6. Eulalia falsa: a head and proboscis; 6 anterior view of parapodium; ¢ seta.
Protomystides capensis: d seta; e anterior end; f anterior view of parapodium.
Syllis trifalcata: g head; h parapodium; i seta.
THE POLYCHAET FAUNA OF SOUTH AFRICA 305
Eulalia trilineata Saint Joseph 1888
Eulalia trilineata. Fauvel 1923, p. 162, fig. 57m.
Eulalia near albopicta Day 1951, p. 20.
Eulalia near trilineata Day 1953, p. 410.
Records: LAM.5(1), 10(1), 22(1), 35(1); FAL.22(2), 27(3), 81(1), 103(2),
113(7), 128(1), 131(1), 145(2), 152(1), 219(1), 245(1), 260(2), 262(2); ©
LIZ.29(1).
Notes: The numerous specimens now available allow me to confirm the
identity of this common South African species with Fauvel’s brief description.
Apart from the colour pattern the setae are quite characteristic. There are
relatively few setae (10—15), the shaft-heads are markedly swollen and the blades
are very short and strongly tapered. In 1951 I suggested that a specimen from
Port St. Johns was close to E. albo-picta Marenzeller from Japan. Since then I
have been able to consult Izuka (1912) who gives an excellent description of
this species. It has tentacular cirrus V, flattened and there are setae on the
second tentacular segment. In F. trilineata V, is almost cylindrical and there
are no setae on the second tentacular segment.
Notophyllum splendens (Schmarda) 1861
Notophyllum splendens. Day 1953, p. 4.08, fig. 2 h-k.
Records: LAM.44(1); TB.302(1), 303(1), 309(1), 310(1); TRA.122(1),
143(1); FAL.27(1), 31(p), 56(1), 80(p), 149(1), 162(1).
Eteone foliosa Quatrefages 1865
Eteone foliosa. Fauvel 1923, p. 174, fig. 62 g-k. Day 1953, p. 411.
Records: SB.183(3), 189(1), 195(1); LB.323(1); TB.go1(1); FB.302(1);
FAL.110(1); FAL.113(3); FAL.228(1); ?FAL.375(2).
Eteone (Mysta) syphodonta (Delle Chiaje) 1822
Eteone (Mysta) siphonodonta. Fauvel 1923, p. 178, fig. 63 e-A.
Records: FAL.349(1); TRA.113(1); SCD.61(1).
Notes: The body is brown to mauve dorsally and pale ventrally. The
prostomium is white, bluntly triangular and depressed with a pair of eyes which
are visible through the skin. The two pairs of antennae are subequal and rather
slender. ‘The tentacular cirri are equal. The first normal segment lacks a dorsal
cirrus but has a small ventral cirrus and a setigerous lobe with several setae.
The dorsal cirri are 1-5-2 times as long as broad and borne on rather long
broad cirrophores. The ventral cirri are bluntly pointed and a little longer than
the blunt setigerous lobes. There are 15-20 setae with fairly long, evenly tapered
blades and shaft-heads which are asymmetrical having one large tooth and
3-5 denticles. The proboscis when dissected proved to have two ventro-lateral
306 ANNALS OF THE SOUTH AFRICAN MUSEUM
rows of large triangular papillae, a broad brownish dorsal band of minute
flattened and denticulate papillae and a much narrower ventral band of
slightly larger globular papillae.
This is the first record from South Africa but the above description agrees
very well with that of Fauvel with the exception that the dorsal cirri are a
little longer.
Eteone sp.
Records: TRA.108(1).
Notes: Specimen ‘TRA.108.K is an unidentified species of Eteone new to
South Africa. It is dirty white in alcohol and 15 mm. long. The prostomium
is unusually long and slender and somewhat reminiscent of a Glycerid. It is
tapered and over twice as long as the basal breadth with two pairs of stumpy
antennae one behind the other and a pair of well-developed eyes posteriorly.
Behind the eyes the head swells out to encompass the proboscis which had been
lost. The two pairs of tentacular cirri are very small; the dorsal pair is no
more than an elongate papilla and the ventral is only one-third the breadth
of the tentacular segment. The next segment has only a ventral cirrus there
being neither a dorsal cirrus nor setigerous lobe nor setae. All the parapodia
are small. The dorsal cirrus is roughly semicircular and no broader than its
cirrophore. The setigerous lobe is rather elongated with a blunt apex and the
ventral cirrus is ovoid. There are about 10 setae per bundle each having an
asymmetrical shaft-head with a large tooth on one side and a minute one
borne on a projecting lobe on the other. The blade is broad basally but tapers
rapidly to a slender tip.
This species is definitely new to South Africa and the slender prostomium
and tiny tentacular cirri suggest that it may be a new species but until the nature
of the proboscis is known it is not advisable to give it a specific name.
Protomystides capensis n. sp.
(Fig. 6 d-f)
Records: TRA.86(1); WCD.28(1).
Description: The holotype is a slender orange worm from TRA.86 richly
speckled with red. It is 17 mm. long by 0-7 mm. wide with 110 segments, and
is well tapered at each end.
The prostomium (fig. 6¢) is small and cordate, a little longer than broad
with two pairs of slender antennae and a pair of laterally placed eyes. Dissection
showed that the proboscis had been lost. There are 3 pairs of small subulate
oS, I I
tentacular cirri on three segments according to the formula 1 + S— + S_.
N
The first tentacular segment is fused to the prostomium and its cirrus is
cylindrical and markedly tapered distally. The second segment is broad and
THE POLYCHAET FAUNA OF SOUTH AFRICA 307
distinct, its dorsal cirrus is oval in section and about 1-5 times as long as the
prostomium. The setigerous lobe is well developed and bears several setae.
The ventral cirrus (V,) is quite definitely similar to those of normal body
segments. The third tentacular segment is also distinct but rather narrow and its
cylindrical dorsal cirrus is markedly tapered and only two-thirds the length
of that on the second segment. The setigerous lobe, setae and ventral cirrus
are similar to those of the succeeding body segments.
Normal body segments are depressed and the dorsal cirri are well to the
sides so that most of the back is uncovered. Each dorsal cirrus (fig. 6f) is
symmetrically cordate and about as broad as long. The setigerous lobe is
rather long and has a simple blunt apex, the presetal lobe not being notched
as is the case in most Phyllodocids. The ventral cirrus is ovoid and possibly a
little longer than the setigerous lobe. There are about 12-18 compound setae
(fig. 6d) with swollen, symmetrical and almost truncate shaft-heads which bear
a series of very fine subequal teeth distally. The blade is short and strongly
tapered.
Protomystides is a rare genus and as far as I am aware no species has been
described from the southern oceans, certainly none has been described from
South Africa. Bergstrém (1914) describes P. bidentata from the North Atlantic
and Mediterranean as having all the tentacular segments fully developed, and
free from the prostomium.
Family HESIONIDAE
Syllidia armata Quatrefages 1865
Magalia perarmata Mar. et Bobr., Fauvel 1923, p. 246, fig. 92.
Records: SB.115(1), 183(3), 184(1), 207(1); TRA.86(1), 88(1); FAL.31(1),
43(2), 136(6), 145(1), 164(2), 174(1), 266(1), 275(1), 283(1); MB.88(1);
LIZ.9(2).
Kefersteinia cirrata (Keferstein) 1863
Kefersteinia cirrata. Fauvel 1923, p. 238, fig. 89 a-e.
Records: FAL.283(4).
Notes: This is a new record for South Africa, but the characters agree
very well with Fauvel’s description.
Family SYLLIDAE
Syllis (Haplosyllis) spongicola Grube 1855
Syllis (Haplosyllis) spongicola. Fauvel 1923, p. 257, fig. 95 a-d.
Records: AFR.707(1), 842(1); TRA.151(1); MB.16(2); SCD.54(1).
308 ANNALS OF THE SOUTH AFRICAN MUSEUM
Syllis (Haplosyllis) trifalcata n. sp.
(Fig. 6 g—z)
Records: FAL.216(1).
Diagnosis: The dorsal cirri have 8-12 joints, and the setae have 3 falcate
teeth.
Description: ‘The holotype is 9 mm. by 0-4 mm. with 88 segments. There
are no colour markings. The head (fig. 6g) is broader than long with rather
flattened palps bent ventrally but not united at the base. The antennae are
subequal and rather short. There are 4 eyes. The pharynx has an anterior
dorsal tooth and extends back to setiger 9 and the cylindrical proventriculus
with 40 rows of points then extends on to setiger 16.
The tentacular cirri and dorsal cirri are short, tapered and twist like
pigs’ tails; they have 9-12 well-marked joints. The setigerous lobes (fig. 6h) are
obliquely truncate cones and the ventral cirri are small. There are 2 acicula
with blunt tips. Each parapodium contains 3-6 simple setae (fig. 62) which are
all similar, each having an expanded end (corresponding to the shaft-head)
bearing 3 claw-like teeth of about the same size.
The common S. (#.) spongicola Grube has dorsal cirri with more joints
and the setae are roughly like boat-hooks with 2 small teeth above a large
triangular rostrum. S. (H.) depressa Augener 1913 from Australia has setae
with only 2 teeth rather like the open beak of a bird. S. (H.) abberans Fauvel
1919 from Indochina is fairly close but the dorsal cirri are long and apparently
not jointed; moreover the setae are narrowed before the apex which has teeth
approaching those of S. (H.) spongicola.
Syllis vittata Grube 1840
Syllis vittata. Fauvel 1923, p. 263, fig. 98 7-1. Day 1953, p. 412.
Records: LAM.22(1); FAL.82(1), 134(1), 171(2); MB.66(r).
Notes: Specimens LAM.22.W and FAL.171.Z are doubtfully referred to
S. vittata. The body is creamy white without markings and rather stout. The
pharynx is short. Dorsal cirri have about 20 joints. The setae always have a
very small secondary tooth and in the middle of the body they tend to be
short and hooked.
Syllis variegata Grube 1860
Syllis variegata. Fauvel 1923, p. 262, fig. 97 h—n. Day 1953, p. 412.
Records: LAM.22(1); SB.189(1), 197(2); WCD.8(2), 19(4); FB.307(4);
FAL.8(p), 113(2), 128(1), 131(1), 134(1), 145(2), 156(5), 162(12), 174(2),
178(2), '302(1),'303(1);) MB.86(1)5 LIZ.2(1) (ny 20K n)).
3
THE POLYCHAET FAUNA OF SOUTH AFRICA 309
Syllis prolifera var. zonata (Haswell) 1886
Syllis zonata Augener 1918, p. 236, pl. 4, fig. 86; pl. 5, fig. 107, text-fig. 19.
Records: LAM.25(1), 31(1), 44(1), 47(1), 57(1), 59(1); SB.197(1);
TB.332(1); FAL.31(8), 81(p), 103(1), 110(p), 128(1), 134(1), 145(2), 149(3),
156(3), 162(20), 166(2), 171(19), 219(1), 275(2), 280(1); LIZ.18(1).
Notes: Like S. variegata this species has dorsal cirri with 25-35 joints and
strongly bidentate setae; it differs in having a short pharynx and in having two
narrow black lines across the anterior segments where S. variegata has a pattern
of broken brown bars.
Syllis armillaris Miller 1776
Syllis armillaris. Fauvel 1923, p. 264, fig. 99 af. Day 1953, p. 412.
Records: Lamberts Bay 13 records from 17-23 metres on rock. Common.
SB.207(1); TB.305(6), 306(1), 308(2), 309(1), 331(4); SH.168(2), 204(1),
366(1), 415(2); WCD.8(2); False Bay—27 records from o—33 metres on rock
(common). AFR.835(1), 967(1); MB.13(1); LIZ.18(1), 36(2), 37(2); SCD.
9(2), 22(1), 106(1).
Syllis gracilis Grube 1840
Spllis gracilis. Fauvel 1923, p. 259, fig. 96 f-i. Day 1953, p. 412.
Records: FAL.17(2), 50(1), 166(2), 275(2), 280(1); MB.86(1); LIZ.2(1).
Syllis hyalina Grube 1863
Syllis capensis McIntosh 1885, p. 193, pl. 33, figs. 8-9; pl. 15A, fig. a1.
Syllis hyalina. Fauvel 1923, p. 262, fig. 98 a—b.
Records: MB.57(1), 86(1).
Notes: The type of Syllis capensis from the Cape is now in the British
Museum. It is a small worm and probably immature. The dorsal cirri are
cylindrical not fusiform and have 13 joints anteriorly, 11 in the middle of the
body and 10 posteriorly. There are 6-10 strongly bidentate setae per foot.
The specimens from Mossel Bay (MB.57 and 86) are referred to S. hyalina
with some hesitation for the dorsal cirri have 15-20 joints. The setae are
strongly bidentate but in superior setae the two teeth are very close together
and project at right angles to the blade somewhat in the fashion shown by
Gravier for S. bouviert.
Syllis cf. trapobanensis Willey 1905
(Fig. 7a)
Typosillis taprobanensis Willey 1905, p. 268, pl. 3, figs. 77, 78.
Records: TRA.55(4); SCD.54(1).
310 ANNALS OF THE SOUTH AFRICAN MUSEUM
Notes: There are faint transverse bars across the anterior segments when
fresh but these soon fade in alcohol leaving the body white. The body is of the
usual size (16 mm.) and shape. The palps are rather flattened and short but
separate at the base. The pharynx is strongly chitinised and the dorsal tooth is
rather small. There is no sign of an occipital flap. The dorsal cirri are rather
long and markedly tapered with 20-30 joints. The setigerous lobes are stout
and the ventral cirri slender. The setae (fig. 7a) are characteristic and all
similar. There are 10-12 per bundle and each has a swollen shaft-head with a
short almost triangular blade with two large blunt teeth.
Syllis (Langerhansia) anops Ehlers 1897
Syllis (Ehlersia) anops Ehlers 1897, p. 40, pl. 2, figs. 40-45.
Records: FAL.248(1).
Notes: A single incomplete specimen was obtained measuring 16 mm. for
45 segments. It is a threadlike worm with slender dorsal cirri. The prostomium
lacks eyes, has large palps fused at the base and short antennae. The tentacular
cirri are a little longer but still shorter than the anterior dorsal cirri. The pharynx
has the dorsal tooth at its anterior margin and stretches back to setiger 9.
The proventriculus is also long and extends back from setiger 9 to setiger 18.
The uniformly slender dorsal cirri are about as long as the body is broad and
have 20-25 joints. The setigerous lobe is in the form of an obliquely truncate
cone and the ventral cirrus is slender and a little longer than the foot. The
setae are similar throughout. In each foot there are 2—3 superior setae with very
long tapering swordlike blades which give the impression of having minutely
knobbed tips. Below these there are about 12 setae with unidentate blades of
normal length.
The South African specimen agrees with Ehlers’ species from the Magellan
area in all respects save one. Ehlers states that the anterior setae are bidentate
and his figure (pl. 2, fig. 44) shows both the Ehlersza type and the normal setae
each with a small secondary tooth. In my specimen all setae agree with his
pl. 2, fig. 45, which shows the blades of both types of setae with unidentate tips.
This is a new record for South Africa.
Syllis (Langerhansia) ferrugina Langerhans 1881
Syllis (Ehlersia) ferrugina. Fauvel 1923, p. 269, fig. 100 k—u.
Records: SB.183(3), 189(1); FAL.149(1).
Notes: This is the first record from South Africa but Augener (1918)
recorded it from Angola.
Syllides longocirrata Oersted 1845
Syllides longocirrata. Fauvel 1923, p. 284, fig. 108 a-g.
Records: FAL.65(1), 82(1).
Notes: The body is small and the pharynx lacks teeth. The antennae,
tentacular cirri and the first two pairs of dorsal cirri are unjointed but the
THE POLYCHAET FAUNA OF SOUTH AFRICA 311
Fic. 7. Syllis cf. taprobanensis: a seta.
Trypanosyllis ankyloseta: b seta; ¢ entire animal; d anterior end; e middle parapodium.
Lamellisyllis comans: f anterior end; g head with left antenna and dorsal cirri removed;
h parapodium; 7 seta.
312 ANNALS OF THE SOUTH AFRICAN MUSEUM
remaining dorsal cirri are very long and deeply annulated with 12-15 joints.
The setae have long tapered blades with indistinct tips which are possibly
unidentate. This is a new record for South Africa.
Trypanosyllis zebra Grube 1860
Trypanosyllis zebra. Fauvel 1923, p. 260, fig. 101 a-e. Day 1953, p. 413.
Records: FB.322(1); LIZ.o(1).
Trypanosyllis gemmulifera Augener 1918
Trypanosyllis gemmulifera Augener 1918, p. 278, pl. 5, figs. g9-101, text-fig. 27. Day 1953, p. 413.
Records: LAM-4(¢), 1o0(1), 15(2), 18(3), 22(3), 33(1)5)@ 9) ayaa
5B.132(2); LB.161(1); TB.306(2); SH.324(1); AFR.842(1); TRAR@os(3)
r10(1); WCD.8(3); FAL.57(p), 103(p), 106(p), 145(1), 140(5)aumy olay.
184(1), 216(2); MB.g(1), 13(1), 16(1), 53(2), 67(3), 77(1), 86(1); LIZ.29(3).
Trypanosyllis prampramensis Augener 1918
Trypanosyllis prampramensis Augener 1918, p. 276, pl. 4, figs. 91, 92, text-fig. 26. Day 1953, p. 414.
Records: FAL.156(1).
Trapanosyllis ankyloseta n. sp.
(Fig. 7 b-e)
Records: FAL.216(1).
Diagnosis : A short broad body, dorsal cirri with 6-8 joints, simple setae with
the blade fused to the shaft-head.
Description: ‘The holotype is the single specimen dredged in False Bay at
32°12-4'5/18°43°5'E at a depth of 42 metres on a sand and rock bottom. The
specimen (fig. 7c) is very broad and short and markedly flattened. It measures
8 mm. by 1°8 mm. and is roughly oval with about 120 segments. The colour
is yellowish white in alcohol.
The prostomium (fig. 7d) which is sunk in between the anterior segments,
is rectangular with 4 large eyes, a pair of ovoid palps directed ventrally and 3
short antennae. The lateral pair are anterior in origin and have 6 joints while
the median arises from the centre of the prostomium and has 8 joints. The
tentacular cirri are borne on anteriorly directed projections arising between
the prostomium and the first pair of parapodia and the tentacular segment is
not visible dorsally. The dorsal pair of tentacular cirri are longer than the
ventral pair and are about equal to the dorsal cirri of setiger 1. The mouth is
ventral and the long pharynx is folded on itself in the dorso-ventral plane. The
trepan has about 10 teeth. The proventriculus which has about 40-50 rows
of points extends from setiger 16-26.
THE POLYCHAET FAUNA OF SOUTH AFRICA 313
Anterior segments increase in width until an average segment in the middle
of the body is about 20 times as broad as long. Posterior ones decrease again
as the body tapers to an oval anterior end. The parapodia (fig. 7e) are similar
throughout. Each has a short dorsal cirrus of 6 to 8 joints borne on a broad
projecting cirrophore. Below this is the setigerous lobe with a terminal papilla
and below this again is the somewhat shorter ventral cirrus. The 4-5 setae
(fig. 7b) are not compound but simple since the falcate unidentate blade has
become fused to the shaft-head. The posterior end tapers to a bilobed pygidium
bearing a pair of ovoid anal cirri with 3-4 joints.
Odontosyllis polycera (Schmarda) 1861
Odontosyllis polycera. Augener 1918, p. 283, pl. 5, fig. 97. Day 1953, p. 415.
Records: LAM.31(2); FAL.23(p), 31(1), 50(3), 82(2), 104(p), 113(2),
164(1), 365(1); MB.57(2), 87(4).
Odontosyllis ctenostoma Claparede 1863
Odontosyllis ctenostoma. Fauvel 1923, p. 277, fig. 104 f-l.
Records: ‘TRA.121(1).
Notes: ‘This species has been recorded from Angola by Augener (1918)
but this is the first record for South Africa.
Pharnygeovalvata natalensis Day 1951
Pharyngeovalvata natalensis Day 1951, p. 26, fig. 4 e-j.
Records: FAL.171(1).
Notes: ‘The single specimen is incomplete but measured 9 mm. for 48
segments and is thus larger than the type. It has a general resemblance to
Odontosyllis ctenostoma but the structure of the pharynx is characteristic.
Amblyosyllis lineolata (Costa) 1864
Pterosyllis formosa Clap. Fauvel 1923, p. 280, fig. 105 h-n.
Amblyosyllis lineolata Day 1953, p. 415.
Records: FAIL.136(2), 159(1), 162(1), 171(1).
Pionosyllis ehlersiaeformis Augener 1913
Pronosyllis ehlersiaeformis Augener 1913, p. 225, pl. 3, fig. 32; text-fig. 31 a-e.
non Pionosyllis ehlersiaeformis Day 1953, p- 415, fig. 3d.
Records: WCD.13(8); FAL.269(4).
Notes: The material consists of four ovigerous females 4-8 mm. long living
in mucus tubes attached to hydroids. Two of them had developing embryos
on their backs.
314 ANNALS OF THE SOUTH AFRICAN MUSEUM
The prostomium bears 3 antennae on its anterior margin and the median,
which is the longest, is twice the length of the prostomium. There are 4 large
eyes and broad palps which are united basally and bent ventrally. The
tentacular segment is short and distinct from the prostomium. It bears two
pairs of tentacular cirri, the longer dorsal pair being equal to the median
antennae.
The dorsal cirri are smooth and tapered. The pair on the first setiger are
about 1°5 times the breadth of the body but succeeding ones are shorter and
over most of the body the dorsal cirri are only two-thirds the breadth of the
body. Some of the cirri are wrinkled but none are annulated.
There are 10-12 setae per foot. Those of the first foot are stouter than the
rest and at certain angles the blade almost appears fused to the shaft but in
the following feet the majority of the setae are clearly compound with bidentate
blades. A single slightly curved, simple needle-like seta appears on the roth
foot and a little later two Ehlersia-type compound setae with very long slender
blades. Beneath these are several compound setae with shorter blades with
two large terminal teeth. An inferior simple seta was not seen but in two
specimens natatory setae with immensely long blades were found in posterior
feet.
Augener’s descriptions of the dorsal cirri are not quite consistent for in the
first account (Augener 1913) he states that the dorsal cirri are not ringed and
in the second (Augener 1918) he says that they are.
The tendency for transverse wrinkling or indistinct ringing is common
in the genus and it is possible that the different descriptions are due to variations
in methods of preservation.
Augener (1918) suggested that P. malmgreni described by McIntosh (1904)
from False Bay as having 20-30 annulations to the dorsal cirri was synonymous
_with his P. ehlerstaeformis. In Day (1953) I followed Augener but noted that my
specimen did not have true Ehlersia-type setae. I now feel that they should be
kept separate for the Cape form of P. malmgreni lacks the Eflersia-type superior
setae, has 20-30 indistinct joints to the dorsal cirri and grows to a much larger
size (40 mm. as against 8 mm.).
Pionosyllis cf. longocirrata St. Joseph 1887
Pionosyllis cf. longocirrata. Fauvel 1923, p. 288, fig. 110 h-l.
Pionosyllis sp. Day 1953, p. 418, figs. 3 ef.
Records: SH.204(2), 415(1).
Notes: These specimens show many similarities to Fauvel’s description but
the identification remains uncertain. All three individuals are soft and fragile
and have broken in several places but might measure 15 mm. if complete.
The prostomium has 2 pairs of eyes and the anterior pair are larger and further
apart. The palps are fused at their bases and bent ventrally. The pharynx is
short with a smooth rim and a fairly large anterior tooth. The proventriculus
THE POLYCHAET FAUNA OF SOUTH AFRICA 315
has about 30 rows of points. The dorsal cirri are very long, smooth, tapered and
over twice the breadth of the stout body. The anterior ventral cirri are large
and triangular but not lamellar and further back they become more digitiform.
The setae are very long and fine. The blades themselves are slender, and not
obviously tapered. The tips are bidentate and as shown in Day (1953) figs. 3 e-f,
there is a strong hooked terminal tooth with a slender tooth directed obliquely
towards it. There are also faint indications of a hood over the terminal tooth.
No simple setae were seen.
The European P. longocirrata has even longer dorsal cirri, up to 4 times the
body breadth but Fauvel’s figure 110h does not suggest that they are tapered
and his figure of the seta (110/) does not suggest a hooked terminal tooth.
Pionosyllis magnidens Day 1953
Pionosyllis magnidens Day 1953, p- 416, fig. 3 a—c.
Records: FAL.132(2), 174(3), 178(1).
Grubea furcelligera Augener 1913
Grubea furcelligera Augener 1913, p. 256, pl. 3, figs. 20, 21; text-figs. 39.
Records: FAL.275(2).
Notes: The material consists of two females with natatory setae carrying
developing embryos on their backs. They are colourless in alcohol. The best
preserved measures 5 mm. in length and has 40 segments. The prostomium is
rounded with a pair of palps which are square in front, fused for most of their
length but with the distal ends free. There are 3 antennae all arising from the
anterior margin of the prostomium. All are bottle-shaped with tapered ends
and the median is twice the length of the laterals which are equal to the width
of the body (not including the parapodia) at the level of setiger 1. There are
4 eyes. The pharynx extends back to setiger 4 with the dorsal tooth near the
anterior end. The barrel-shaped proventriculus extends over a further 2
segments.
The tentacular segment is clearly marked off from the prostomium but is
very narrow and not very distinct from setiger 1. It bears the usual 2 pairs of
tentacular cirri, of the same elongate subulate shape as the antennae. The dorsal
pair which is twice as long as the ventral pair is equal to the median antenna.
The dorsal cirri are similar to the antennae and the tentacular cirri but vary
in length; the dorsal cirri of setiger 1 are ? the length of the antennae, those of
setiger 2 are very short and hardly exceed the length of the setigerous lobe,
those of setigers 3 and 4 increase again and the dorsal cirri of setigers 5 and
subsequent segments are about half as long as the tentacular cirri or a little
over half the width of the segment that bears them. The setigerous lobes are
short truncate cones and the ventral cirri are rather stout and ovoid.
The normal compound setae have swollen shaft-heads and very small,
unidentate dagger-like blades. There is also a single pointed simple seta in the
316 ANNALS OF THE SOUTH AFRICAN MUSEUM
superior part of each bundle. The natatory setae appear in setiger 9. Each
has a long slender shaft from which arises a very fine hair-like tapered blade.
This South African material lacks the small anterior third pair of eyes
described by Augener and the antennae, tentacular cirri and dorsal cirri of
setiger 1 are considerably longer than Augener describes, possibly due to
differences in preservation. This is a new record for South Africa.
Grubea rhopalophora Ehlers 1897
Grubea rhopalophora Ehlers 1897, p. 53, pl. 3, figs. 66-70. Augener 1918, p. 295, pl. 4, fig. 94.
Records: FAL.17(1), 246(2); MB.85(1).
Notes: The Cape material agrees perfectly with Ehler’s description and I
can confirm that the compound setae are unidentate. ‘This species has previously
been recorded by Augener (1918) from the shore at Swakopmund and shallow
dredgings at Liideritzbucht in South West Africa. In his notes on individuals
from Liideritzbucht Augener describes two with minute and truncate dorsal
cirri containing fibrillar structures. These agree very closely with the description
of Fauvel (1923) of Grubea pusilla (Dujardin).
Grubea rhopalophora is generally similar to G. limbata Claparéde but differs
in the fact that the antennae, and dorsal cirri are shorter with more swollen
bases, the tentacular segment is more completely fused with the prostomium
and the palps separate towards their extremities.
Sphaerosyllis sublaevis Ehlers 1913
Sphaerosyllis sublaevis Ehlers 1913, p. 482, pl. 32, figs. 10-15.
Records: SB.167(1); FAL.82(1); TRA.113(1).
Notes: These specimens agree very well with Ehlers’ description. The
body surface is smooth and there is no dorsal cirrus on setiger 2. ‘The antennae,
tentacular cirri and anterior dorsal cirri are all small and flask-shaped with
swollen bases and tapered ends but further back the dorsal cirri become longer
and more bottle-shaped. ‘The setae have unidentate blades. Ehlers states that
there is a third minute pair of eyes on the anterior margin of the prostomium.
Their absence in the present specimens is not regarded as important as it has
been noted that such eye-specks are often invisible in individual specimens of
Syllids.
S. sublaevis is close to S. claparedit Ehlers (1864) but the latter is reported
to have a dorsal cirrus on setiger 2.
Sphaerosyllis hystrix Clap. var. capensis Day 1953
Sphaerosyllis hystrix Day 1953, p. 420, fig. 4 g-l.
Records: SB.183(2).
THE POLYCHAET FAUNA OF SOUTH AFRICA 317
Exogone clavator Ehlers 1913
Exogone clavator Ehlers 1913, p. 485, pl. 33, figs. 1-6. Day 1953, p. 418.
meoras =) WOOD .5(1) >, 5b.183(4); FAL.17(7), 110(1),. 131(6), 152(1),
159(2), 266(1); MB.57(1); LIZ.20(1).
Exogone gemmifera (Pagenstecher) 1862
Exogone gemmifera. Fauvel 1923, p. 305, fig. 117 a-d.
Exogone verugera (non Claparéde) Day 1953, p. 418.
moms oh.107( 2); FAL:29(7), 82(2), 103(3), 128(1), 178(10), 280(1).
Notes: In 1953 I referred several specimens to E£. verugera though it was
noted that they lacked a dorsal cirrus on setiger 2. Examination of a great deal
more material from both South Africa and Europe and the discovery of the
typical E. verugera in South Africa has shown that the presence or absence of a
dorsal cirrus on setiger 2 is a constant and important character. It is now possible
to summarize the main differences between the three closely related species
E. verugera, E. gemmifera and FE. heterosetosa, all of which occur in the Southern
hemisphere.
In £. gemmifera the palps are short and broad, the three antennae are of
equal size and about the same length as, or a little longer than, the prostomium.
The proventriculus is short, extending over 1-2 segments, and has 10-12 rows of
points. ‘he superior compound seta has a long dagger-like blade. There is no
dorsal cirrus on setiger 2. In E. heterosetosa the palps are short and the three
antennae are about the same length as the prostomium. The proventriculus
is rather long, extending over 3 segments and has 15 or more rows of points.
The superior compound seta has a characteristically swollen shafthead and a
short broad blade. There is no dorsal cirrus on setiger 2. In E. verugera the
palps are rather long and tapered and the three antennae are equally minute
and much shorter than the prostomium. The proventriculus is fairly long,
extending over 2—3 segments and has 25-30 rows of points. The superior seta
has a dagger-like blade. There is a dorsal cirrus on setiger 2.
Exogone verugera Claparéde 1868
Exogone verugera. Fauvel 1923, p. 307, fig. 117, figs. m—r.
Records: SH.400(1); FAL.162(13); SCE.54(1).
Notes: The diagnostic characters of this species are given above.
Autolytus charcott Gravier 1906
Autolytus charcoti Gravier 1906, p. 7, pl. 1, figs. 1, 2.
? Autolytus afer Ehlers, 1908b, p. 46.
iecords.; VB.30e(1yinFAL.o497(1)5(LIZ.29(1).
318 ANNALS OF THE SOUTH AFRICAN MUSEUM
Notes: Specimen TB.312 measures 15 mm. for 75 segments. The body has
conspicuous black bands at the intersegmental junctions starting at setiger 1/2
then every junction to setiger 6/7, then misses 7/8 and 14/15 and thereafter
is present on every fourth junction to the end of the body. On specimen
LIZ.29 every intersegmental junction from 2/3 onwards is banded.
Diverging nuchal epaulettes extend from the back of the prostomium to
setiger 2. The trepan has 10 equal teeth. The antennae, tentacular cirri and
the dorsal cirri of setiger 1 are stout and just longer than the width of the body,
but thereafter the dorsal cirri decrease and in the middle of the body they are
rather less than one-third the body width.
The specimen described by Ehlers (1908) from Liideritzbucht as A. afer
agrees in general characters, but as it was preserved in osmic acid no details
of colour pattern are available.
Autolytus tuberculatus (Schmarda) 1861
Autolytus tuberculatus. Augener 1918, p. 307. Day 1953, p. 421.
Records: FAL.22(1), 82(7), 103(3),.113(1), 122(1), 145(2); 150( pees
171(2), 247(1), 290(5) 3-2 LIZ.56(1):
Notes: Further material has shown that the length of the nuchal epaulettes
is variable. They may reach setiger 6 or hardly reach setiger 4. Anterior dorsal
cirri are unequal; those of setigers 1, 2, 4 and 6 are much longer than those of
setigers 3, 5 and 7 or subsequent segments. In the middle of the body they are
only one-third to one-half of the width of the body.
Autolytus prolifer (Miller) 1788
Autolytus prolifer. Fauvel 1923, p. 311, fig. 119.
Records: FAL.334(1); MB.58(1), 69(1); SCD.40(1).
Notes: The pharynx is S-shaped and crowned with 10 large triangular
teeth. Indistinct nuchal epaulettes are present on setiger 1 but do not extend
on to setiger 2. The antennae, tentacular cirri and dorsal cirri of the first
setigers are long but the remaining ones are only one-third to one-quarter the
width of the body.
Autolytus maclearanus McIntosh 1885
Autolytus maclearanus. Ehlers 1913, p. 488, pl. 33, figs. g—-11.
Autolytus inermis (non St. Joseph) Ehlers 1913, p. 488.
Records: SH.430(20); FAL.43(1); SCD.61(3).
Notes: This South African material agrees very well with the species
described by Ehlers from Kerguelen under the name of A. maclearanus.
McIntosh’s original description is so vague that it might refer to any species of
Autolytus, but Ehlers’s description and figures are clear.
THE POLYCHAET FAUNA OF SOUTH AFRICA 319
The diagnostic features are the long antennae, dorsal tentacular cirri and
dorsal cirri of setiger 1. All of these greatly exceed the width of the body and
are often so wrinkled as to give the impression of being annulated which they
are not. The dorsal cirri of normal body segments are about one-third of the
body width. The tentacular segment is much shorter than setiger 1 and on some
specimens it has vague indications of small nuchal epaulettes as stated by Ehlers.
The pharynx has 6 rounded lappets instead of sharp chitinous teeth at its
entrance. These were only seen when the pharyngeal sheath was dissected
away and the pharynx was first thought to be unarmed as in A. inermis. The
latter species, however, has a doubly convoluted pharynx while the present
species has a single large loop.
Ehlers stated that the proventriculus of his Kerguelen specimen lay in the
7th segment. Here it is in the 4th and has 30 rows of points. In fresh material
it is faintly greenish.
Myrianda phyllocera Augener 1918
Myrianida phyllocera Augener 1918, p. 301, pl. 4 figs. 87-89, text-fig. 30. Day 1953, p. 421.
Records: FAL.178(1); LIZ.2(1).
Lamellisyliis gen. nov.
Prostomium with 3 lamellar antennae. Palps united at their bases.
Pharynx straight with an anterior dorsal tooth. Prominent nuchal epaulettes.
A single pair of cylindrical tentacular cirri. Dorsal cirri lamellar, setae
compound, ventral cirri on all segments. Type species L. comans.
Lamellisyllis comans n. sp.
(Fig. 7-1)
Records: FAL.110(1).
Description: The holotype is a pale, flattened worm, roughly Harmothoid
in outline and measures 8 mm. for 50 segments. The prostomium is sunk back
between the anterior segments which project forwards and outwards so that
the front end of the body (fig. 7/) appears rounded. The palps are normal and
united only at their bases. The small rounded prostomium (fig. 7g) has 4 eyes
set in a rectangle and 3 subequal foliaceous antennae. The lateral pair arise
from the anterior margin while the median arises from the centre of the prosto-
mium. The mouth is ventral and the straight, weakly chitinized pharynx
bears a single dorsal tooth near its anterior margin. It extends back to setiger 7
and the barrel-shaped proventriculus with 20 rows of points extends on to
setiger 12.
‘T'wo grooved, finger-like nuchal organs diverge from the posterior margin
of the prostomium towards the sides of setiger 3. They were first thought to
lie freely on the dorsum but attempts to move them showed that they are
320 ANNALS OF THE SOUTH AFRICAN MUSEUM
attached throughout their length. A single pair of tapered and cylindrical
tentacular cirri project forwards on either side of the prostomium. This pair
corresponds to the ventral cirri of a normal segment for each arises from a lobe
of the tentacular segment which is wedged between the prostomium and the
first setiger and above it there is a lump which seems to correspond to a dorsal
cirrophore.
The normal body segments (fig. 7h) are all similar. Each is about 20 times
as broad as long and has a dorsal cirrus, a setigerous lobe and a ventral cirrus
on its lateral margin. The dorsal cirrus is borne on a stumpy cirrophore placed
well above the setigerous lobe and the cirrus itself is flattened, oval to circular
in outline, and is attached to the cirrophore by its edge. Alternate dorsal cirri
are more medial and more lateral in origin. ‘The setigerous lobe is an obliquely
truncate cone with a minute papilla at its apex. There is a pointed aciculum
and about 20 compound setae (fig. 77) whose blades are strongly bidentate and
‘hairy’. The ventral cirrus is conical and slightly shorter than the setigerous
lobe. The posterior end of the worm is markedly tapered and the pygidial
segment bears a pair of foliaceous anal cirri.
The possession of foliaceous appendages is unusual in the family Syllidae.
The genus Myrianida has flattened head appendages and dorsal cirri but the
completely fused palps, the sinuous pharynx crowned with a trepan of teeth,
the minutely-bladed compound setae and lack of ventral cirri immediately place
it in the sub-family Autolytinae. Phyllosyllzs Ehlers (1897) from South Georgia
obviously belongs to the Autolytinae as well although it has setae on the tentacular
segment. Knox (1957) has recently described Clavisyllis from New Zealand with
inflated, ovoid dorsal cirri, normal ventral cirri, large nuchal epaulettes, palps
united only at their bases and a straight pharynx with an anterior tooth. In
these characters it is very similar to the present Lamellisyllis, but it differs in
having cylindrical antennae, a nuchal cirrus between the nuchal epaulettes
and two pairs of tentacular cirri. Possibly Clavisyllis should be included in the
sub-family Eusyllinae. Lamellisyllis with its single pair of tentacular cirri shows
resemblances to Sphaerosyllis, Exogone and Spermosyllis all of which belong to the
sub-family Exogoninae. But all the members of the latter sub-family, have palps
which are fused throughout their length so it might be better to place
Lamellisyllis in a sub-family of its own.
The characters of Clavisyllis and Lamellisyllis show that the family Syllidae
is closer to the family Phyllodocidae than had previously been realized.
Procerastea perriert Gravier 1900
Procerastea perriert. Fauvel 1923, p. 327, fig. 126 a-c.
Records : SH.430(4); SB.167(LC).
Notes: ‘This is a new record for South Africa but the specimens agree
perfectly with Fauvel’s description. This species seems to feed on hydroids
growing just below low-tide mark.
ae a ee
THE POLYCHAET FAUNA OF SOUTH AFRICA 321
Family NEREIDAE
Laeonereis ankyloseta Day 1957
Laeonereis ankyloseta Day 1957, p. 83, fig. 5 a-y.
Records : FB.302(3), 307(6); FAL.8(1), 43(1), 50(1), 80(p), 110(2), 126(1
I71(1), 225(4), 249(1), 275(1), 304(1), 345(1); MB.13(1), 16(1), a
56(heteronereid), 62(1 juvenile), 86(1), 87(1), 88(3); LIZ.1(1), 9(1); 8
32(3), 54(2)-
Notes: MB.62 is a juvenile which lacks the characteristic ankylosed setae
but the other characters are typical. MB.56 is a heteronereid.
b)
)
dh
Nereis (Neanthes) operta Stimpson 1855
Nereis (Neanthes) operta. Day 1934, p. 38, fig. 5. Day 1951, p. 28. Day 1953, p. 424.
Records: LAM.4(1), 8(1), 16(1), 22(2), 25(1), 44(t), 51(1), 59(1)s 6t(x),
63(1); SB.118(1), 175(2), 179(1), 181(1), 189(15); SH.366(1), 428(1), TB.
302(1), 325(1), WCD.23(1); TRA.69(2 planktonic heteronereids), False Bay:
24 records from 2-38 metres on sand and rock. MB.49(1), 62(1), 87(6), 88(1) ;
LIZ.6(6).
Nereis (Neanthes) willeyi Day 1934
Nereis (Neanthes) willeyi Day 1934, p. 38, fig. 6 a—c. Day 1951, p. 28. Day 1953, p. 424.
Records : False Bay: 22 records from o—22 metres on sandy rocks. MB.40(4),
49(t), 59(1), ?71(1).
Nereis (Neanthes) cf. kerguelensis McIntosh 1885
Nereis kerguelensis. Ehlers 1897, p. 65, pl. 4, figs. 81-93.
Records: AFR.g50(1); 994(1); TRA.143(1).
Notes: ‘The largest of the three specimens measures 22 mm. It is pale
brown in alcohol with a more intense bar across setiger 2. The proboscis has
group I = o; II = a wedge of 8-9 points; III = 5-6; 1V = a wedge of 10
points; V = 0; VI = 2-5 in a close set group; VII and VIII = a single row
of 3-5. Anterior feet have three notopodial lobes and a rather longer dorsal
cirrus. In posterior feet there are only two notopodial lobes. There are no
notopodial falcigers. The neuropodial falcigers have straight blades with a
tendon towards the tip. This South African material differs from published
descriptions of WV. keurguelensis in having more denticles on group VI. In this
and in the brown bar on setiger 2 it resembles WV. unifasciata Willey but the
latter only has two notopodial lobes on anterior feet.
Nereis (Neanthes) succinea Frey & Lueckart 1847
Nereis glandulosa Ehlers 1908a, p. 74, pl. 8, figs. 1-6.
Records: TRA.91(3); LIZ.1(7), 3(10), 38(1).
322 ANNALS OF THE SOUTH AFRICAN MUSEUM
Nereis (Nereis) lamellosa Ehlers 1868
Nereis lamellosa Ehlers 1868, p. 564, pl. 22, figs. 10-17. Fauvel 1936, p. 36.
Records: TRA.33(3), .91(1); FB.306(6); FAL.184(1), 167(@) Seema:
209(1), 223(4), 240(2); MB.4(3), 34(2), 75(2); SCD.20(1), 61(1), 94(3),
105(1).
Notes: As shown by Fauvel (1936) this species is very close to WV. succinea
but has dorsal homogomph falcigers in posterior feet. All neuropodial falcigers
have feathered blades with a tendon from the apical tooth.
Nereis (Nereis) zonata var. persica Fauvel 1911
Neireis zonata var. persica Fauvel 1911, p. 385, pl. 19, figs. 10-16, 18-23; pl. 20, figs. 24-25.
Records: SH.71(1); SCD.50(1), 63(1).
Notes: This species which is well known from the tropical Indian Ocean
has been found as far south as Mogambique (Day 1957). The present records
show that it extends even further south in dredgings off the eastern Cape
Province. It has also been found on the hull of a ship visiting Table Bay from
the Indian Ocean.
Nereis (Nereis) jackson’ Kinberg 1866
Nereis (Nereis) jackson. Fauvel 1932, p. 97.
Records: L1Z.27(2); SCD.100(1).
Nereis (Nereis) falsa Quatrefages 1865
Nereis falsa. Fauvel 1923, p. 337, fig. 129 e—m.
Neries callaoana (non Grube) Augener 1918, p. 174 (partim).
Records: SH.71(1).
Notes: ‘The specimen is typical. Though known from the Natal coast it has
not been recorded from the Cape. The present record is from the hull of a ship
from India. Augener’s 4 specimens from Swakopmund labelled WV. callaoana
(V.8782) were kindly sent to me by the Director of the Hamburg Museum.
Three proved to be WV. falsa and one Platynereis dumerilit.
Nereis (Nereis) eugeniae (Kinberg) 1866
Nereis eugeniae. Ehlers 1897, p. 67, pl. 4, figs. 94-105. Monro 1936, p. 136.
Records: TRA.74(1), 80(3).
Notes: The anterior part of the prostomium is free from the bases of the
palps; the eyes are small and the tentacles are short. ‘The dental formula of the
largest specimen is: I is 0; II is 3-4 in a line; III is 0; IV is 8-9 in a single to
double row; V is 0; VI is a transverse group of 3; VII and VIII consist of 4
THE POLYCHAET FAUNA OF SOUTH AFRICA 323
widely separated points in a single row. All paragnaths are minute and as
shown they are not numerous in these small specimens.
Anterior feet have two dorsal lobes. In the posterior feet all lobes are
pointed and the dorsal lobe is much larger than the rest. By contrast the dorsal
cirrus is thin and delicate. Most posterior notopodia include 1-2 stout homo-
gomph falcigers with very short conical blades hardly longer than the width
of the shaft. Ventral falcigers of anterior feet have long straight blades; in
posterior feet there are fewer but larger ones.
Nicon eugeniae was first described by Kinberg from a specimen collected
off Argentina opposite the Rio de la Plata. Most workers however refer
to the description given by Ehlers (1897) for Nereis eugeniae based on specimens
collected in the Magellan area and compared with Kinberg’s type. It is
significant that neither author refers to homogomph falcigers in the posterior
notopodia, and that Ehlers describes and figures the tentacular cirri of his
specimen as being annulated and says that the tentacular cirri of Kinberg’s
specimen were ‘gegen die Spitze hin deutlich gegliedert’.
I have examined specimens in the British Museum identified by Monro
(1930) and (1936) as Nereis eugeniae. None of these show annulated tentacular
cirri and all have homogomph falcigers with short conical apices in the posterior
notopodia. They are identical with my South African material. Kinberg’s
types must be examined before the identity of Monro’s specimens and mine
can be firmly established.
NV. eugeniae is closely related to WN. trifasciata Grube 1878 from the Phillipines.
In the original description there was again no reference to homogomph falcigers
in posterior notopodia but Augener (1922) who states that he was not able to
see Grube’s type, describes WN. trifasciata from Juan Fernandez Island as having
notopodial falcigers in posterior feet with long, almost straight blades some 4-5
times as long as the width of the shaft. Fauvel (1932) refers to similar setae in
his description of material from the Indian Ocean. Here again the type must
be examined before the naming is certain, but it will be obvious that the main
difference between WN. eugeniae (sensu Monro 1936) and WN. trifasciata (sensu
Augener 1922) lies in the structure of the notopodial falcigers.
Nereis (Nereis) sp.
(Fig. 8a)
Records: TRA.62(1).
Notes : The material consists of a single complete specimen 8 mm. long with
40 segments. The prostomium is broadly rounded in front and the palps are
large with small palpostyles; the tentacles are normal and rather short. There
are no colour markings. The dental formula is I = 1; II is a wedge of close-set
points; III is a few scattered points; IV is like II; V = 0; VI is a rosette of
8-10 points; VII and VIII is a continuous band consisting of 2-3 irregular
rows.
324 ANNALS OF THE SOUTH AFRICAN MUSEUM
Anterior feet have two notopodial lobes. In posterior feet the superior
notopodial lobe is enlarged and flattened and carries the cirrus at its apex;
the inferior notopodial lobe is a slender cone.
Anterior notosetae are homogomph spinigers with short blades. Anterior
neurosetae include homogomph spinigers with short blades and heterogomph
spinigers. In middle feet there is a gradual change in the setae; the notopodial
spinigers decrease in number and some of the neuropodial falcigers lose the
articulation between shaft and blade. In posterior feet the notosetae include
2-3 homogomph falcigers with long straight blades and the neuropodial setae
include 1-2 stout simple hooks (fig. 8a) formed by fusion of the straight pointed
blade with the shaft, 2-3 normal heterogomph falcigers with fairly straight
blades and 1-2 fine homogomph spinigers.
As far as I can ascertain no species with similar setae has been described
but the single specimen is a juvenile of 8 mm. and for this reason is not named
as a new species.
Nereis (Neanthes) caudata (Delle Chiaje) 1841
? Nereis cricognatha Ehlers. Augener 1913, p. 163. Knox 1951, p. 217, pl. 45, figs. 6-8.
Nereis (Neanthes) caudata. Fauvel 1923, p. 347, fig. 135 a-e. Day 1953, p. 425.
Records: LB.161(1); SH.204(1), 366(1); MB.16(4).
Notes: After studying the descriptions of Augener (1913) and Knox (1951)
I see no reason why N. cricognatha Ehlers (1905) from New Zealand should not
be included in the synonymy of N. caudata.
Perinereis capensis (Kinberg) 1865
Perinereis capensis. Monro 1933, p. 495, figs. 7-11. Day 1934, p. 42, figs. 8 a-e.
Records: SB.177 (1 juvenile); FAL.80(p), 122(p), 127(1), 159(2), 171(2),
174(1), 245(1); MB.o(1), 13(1), 16(2), 20(2), 40(4), 49(21), 53(18), 56(2),
62(1), 67(2), 74(2), 78(1), 85(1); LIZ.1(1), 6(4), 18(fc), 27(4).
Pseudonereis variegata (Grube) 1856
Nereis (Mastigonereis) variegata McIntosh 1904, p. 37, pl. 1, figs. 6-10.
| Records: FAL.126(1); TRA.69 (1 swimming in plankton).
Notes: ‘This species which is so common between the tide marks has only
been found on one occasion below low water.
Platynereis dumerili (Aud. and M.-E.) 1833
Platynereis dumerilii. Fauvel 1923, p. 359, fig. 141 a—f. Day 1953, p. 429.
Records: Lamberts Bay 14 records from 10-23 metres on rock (common) ;
SB.114(c), 116(c), 122(p), 127(2), 129(3), 136(1), 180(8), 181(2), 183(3),
184(2), 194(4), 195(c); LB.155(c), 160(5), 161(4), 190(2), 380(4); SH.74(1);
4
THE POLYCHAET FAUNA OF SOUTH AFRICA 325
k
Fic. 8. Nereis sp.: a simple seta from posterior neuropodium.
Glycera benguellana: b jaw support; c papilla from proboscis; d head of compound seta; e
posterior view of foot.
Glycinde capensis: f4—> proboscidial papillae (f} plan view of row /, f?-> lateral view of
rows 2-5); g anterior view of anterior parapodium; A anterior view of posterior parapodium.
Glycinde kameruniana: j4—*> proboscidial papillae (j1 plan view of row, /, 72-5 lateral view of
rows 2—5); k notoseta of posterior foot; / anterior view of anterior foot; m anterior view of
posterior foot.
326 ANNALS OF THE SOUTH AFRICAN MUSEUM
TRA.69 (abundant in plankton); TRA.107(1); False Bay: 33 records from
o-36 metres on Algae or hydroids; (MB.4(1), 20(2), 27(4), 38(1), 40(6),
56(8), 50(1), 74(13), 86(2), 87(2); LIZ 1(2), 6(6), Watr)ieaiay:
SCD.20(1). :
Platynereis australis (Schmarda) 1861
Platynereis magalhaensis Kinberg. Fauvel 1916, p. 434, pl. 8, figs. 21, 22.
Platynereis australis Day 1953, p. 429.
Records: SB.130(7); LB.155(1), 472(3); SH.366(1); 415(1).
Platynereis calodonta Kinberg 1866
Platynereis hewitti Day 1934, p. 44, fig. 9 af.
Platynereis calodonta Day 1953, Pp. 429.
Records: FAL.134(1); MB.40(1); LIZ.6(1), 27(1).
Family SPHAERODORIDAE
Ephesia gracilis Rathke 1843
Ephesia gracilis. Fauvel 1923, p. 377, fig. 148 a—f.
Records: LAM.8(2), 31(1), 54(1), 59(2); SB.119(1), 183(1); TB.305(1),
330(1); TRA.58(1), ?102(3), 143(2); FAL. 223(1), 371(1), SCD.54(1).
Notes: This is a new record for South Africa and the specimens have been
checked as identical with European material.
Family NEPHTHYDIDAE
Nephthys (Nephthys) capensis Day 1953
Nephthys (Nephthys) capensis Day 1953, p. 431, text fig. 5 g—m.
Records: LAM.52(3); LB.364(5); FAL.107(1), 209(1); TRA.104(1).
Notes: In these larger dredged specimens the gills are often cirriform
throughout so that there is a strong superficial resemblance to WV. hombergi.
However, it lacks a bilobed presetal lamella having at most a rudimentary
presetal lamella in the notopodium, and always has short, saw-edged geniculate
setae in the posterior row as well as the usual elongate capillaries.
This species is close to WV. graviert Augener (1913) from Fremantle, and
may indeed be identical. However, Augener’s figures of the anterior setae,
the shape of the parapodial lamellae and the gill suggest that there are important
differences. Moreover, he only mentions two types of setae and gives the
impression that there are only saw-edged geniculate setae in the posterior
row.
THE POLYCHAET FAUNA OF SOUTH AFRICA 327
Nephthys (Nephthys) ? paradoxa Malmgren 1874. .
Records: AFR.1578(1).
Notes: The material consists of the anterior half of a large worm. The
prostomium is pentagonal with 4 subequal antennae and a pair of colourless
eyes. The proboscis has 22 rows of papillae with 5-6 papillae per row. The
ventral cirrus of setiger 1 is a little shorter than the antennae and the dorsal
cirrus shorter still. Branchiae first appear on setiger 9 as foliaceous organs
but become stout and cirriform towards the end of the fragment.
In anterior feet the notopodium has an oval setigerous lobe, a rudimentary
presetal lamella and a rounded posterior lamella a little larger than the
setigerous lobe. The gill is flattened and bears a small dorsal cirrus. The neuro-
podium has a bluntly conical setigerous lobe, a rudimentary presetal lamella
and a rounded postsetal lamella only slightly larger than the feet. The ventral
cirrus is normal. Later feet are essentially similar with slightly more divergent
and pointed rami.
Anterior setae are normal laddered capillaries and posterior setae are long
and bear transverse bands of conical teeth. These are deciduous and where
they have dropped off transverse marks are left.
More material is required before this determination can be confirmed.
Nephthys (Nephthys) hombergu Savigny 1818
Nephthys hombergii. Fauvel 1923, p. 367, fig. 143 a-d. Day 1953, p. 431.
Records: SB.184(3), 202(8), 203(13), 205(1); LB.3o0o(c), 391(2); TRA.
85(1); TB.gor(1); WCD.15(6), 19(3), 21(7), 23(1), 28(6); FAL.187(2),
341(5), SCD.25(1), 63(8), 109(1).
Nephthys (Nephthys) tulearensis Fauvel 1919
Nephthys tulearensis Fauvel 1919, p. 422, pl. 16, figs. 31-39.
Records: MB.81(1).
Nephthys (Micronephthys) sphaerocirrata Wesenberg-Lund 1949
Nephthys sphaerocirrata Wesenberg-Lund 1949, p. 294, figs. 24-26. Day 1953, p. 431.
Records: SB.175(1), 199(2), 202(4), 203(16), 205(2); LB.323(1); WCD.
Hato). 49(20), 21(38), 23(18), 26(14), 28(12); FB.3916(2); FAL.229(1),
250(2), 328(3), 338(1), 352(3), 375(7), 376(3), 378(3); L1Z.25(2) ; SCD.82(2),
109(3).
Nephthys (Aglaophamus) macroura Schmarda 1861
Aglaophamus macroura Hartman 1950, p. 118.
Records: AFR.783(1); AFR.1578(1); SCD.9(4).
Notes: ‘These specimens agree very well with Hartman’s description with
the exception that the branchiae start on setiger 2 not 3 or 4 as stated. It might
328 ANNALS OF THE SOUTH AFRICAN MUSEUM
also be added that the presetal lamella (not described by Hartman) is bilobed.
The lamella is well developed in the first few feet but soon becomes reduced.
It disappears in the notopodia but in the neuropodia minute lobes persist
above and below the setigerous lobe.
Family GLYCERIDAE
Glycera convoluta Keferstein 1862
Glycera convoluta. Fauvel 1923, p. 383, fig. 150 h—n.
Records: LAM.45(2), 49(1), 52(4); SB.125(8), 175(1), 177(14), 179(1),
183(1), 184(2), 189(6), 193(1); LB.158(6), 159(10), 160(2), 169(1), 239(5),
299(c), 300(c), 363(4), 364(6), 380(c), 382(c), 391(6), TB.go1(1); AFR.
1224(1); TRA.77(c), 143(1); WCD.15(5), 19(4), 21(4), 23(5), 26(1), 28(9);
FB.323(7), 330(1); FAL.58(1), 187(1), 205(1), 209(1), 219(1), 225(1), 226(r),
240(9), 241(1), 243(1), 338(1), 341(c), 345(3), 347(2), 376(1), 378(2); KNY.
61(1); LIZ.24(1).
Glycera alba Miller 1788
Glycera alba. Fauvel 1923, p. 385, fig. 150.
Records: ‘TRA.108(2).
Notes: The first gill is on setiger 33. I doubt very much whether this
species is really separate from G. convoluta.
Glycera parashadi Fauvel 1932
Glycera parashadi Fauvel 1932, p. 126, pl. 5, figs. 1-8. Day 1957, p. 86.
Records; FB.317(1); FAL.211(1), 245(1).
Glycera unicornis Savigny 1818
Glycera unicornis. Fauvel 1923, p. 389, fig. 153 e-7. Day 1953, p. 430.
Records: AFR.935(1); TRA.27(1), 91(1); FB.323(3); FAL.117(1), 229(1),
341(1); LIZ.24(1); SCD.18(1).
Glycera papillosa Grube 1857
Glycera papillosa. Kinberg 1857-1910, p. 58, pl. 21, fig. 3. Augener 1922, p. 203, text-fig. 9 a—c.
?Glycera kerguelensis McIntosh 1885, p. 344, pl. 35A, figs. 3-4.
Records: FB.307(1); MB.74(1); LIZ.25(3), 38(1).
Notes : The prostomium has about 8 rings; the proboscidial papillae include
a few ovoid forms, but the majority are slenderly conical, being five times
as long as the basal breadth; they are not ringed. The jaw supports are deeply
forked, the shorter limb being almost half the length of the longer one. There
are no gills. The superior presetal lobe is minute but the inferior presetal lobe
> ee
THE POLYCHAET FAUNA OF SOUTH AFRICA 329
_ is large and pointed. There is a single, broadly rounded post-setal lobe which
reaches the same height as the ventral cirrus.
This species has longer proboscidial papillae and a much smaller superior
presetal lobe than G. capitata.
It has been suggested that Glycera kerguelensis McIntosh 1885 is identical
with G. papillosa and as Augener (1922) says, the description and figures of the
proboscidial papillae agree very well. The type now in the British Museum has
had its head cut off but the remainder of the worm shows that the original was
much larger than Augener’s specimen or mine. In the feet the superior presetal
lobe is much smaller than the inferior one but not quite so minute as in the
present specimens. Otherwise the feet are very alike. G. kerguelensis is certainly
much closer to G. papillosa (type locality Valparaiso, Chile) than it is to
G. capitata (type locality Denmark).
Glycera benguellana Augener 1931
(Fig. 8 b-e)
Glycera capitata v. benguellana Augener 1931, p. 303, text-fig. g.
Records: LAM.26(2), 35(2), 40(2), 41(3), 49(1), 52(1); TB.304(1),
322(1); FAL.51(1), 63(1), 65(1), 206(1), 209(1), 233(3), 250(1); 349(1),
365(1), 378(1); MB.62(1); SCD.109(1).
Notes: The body is tapered at both ends, slightly swollen anteriorly and
the segments are biannulate. The prostomium is very long with numerous ( ?30)
indistinct rings. There are no visible eyes. The jaw supports (fig. 85) have only
one fork developed, the other limb being reduced to a mere expansion of the
base. The proboscidial papillae include numerous elongate forms (fig. 8c)
with 14—16 distinct V-shaped marks on one side and a few stout forms which are
essentially similar in structure. The parapodium (fig. 8e) has a fairly large
dorsal cirrus, two subequal presetal lobes, a single low rounded postsetal lobe
and a broad pointed ventral cirrus. There are no gills.
As shown above this species is quite distinct from G. capitata in the shape
of the jaw supports, proboscidial papillae and even the feet for the superior
presetal lobe of G. capitata is distinctly smaller than the inferior one. Augener’s
record was from deep water off South West Africa but the present records
show that it is common all round the Cape.
Glycera longipinnts Grube 1878
Glycera longipinnis. Fauvel 1932, p. 125, pl. 4, figs. 11-14.
Records: FAL.211(1).
Notes: The prostomium has about 12 poorly marked rings and rather long
terminal antennae. There are no visible eyes. The proboscidial papillae are
elongate and conical but not ringed. Gills start on the 2oth setiger as simple
330 ANNALS OF THE SOUTH AFRICAN MUSEUM
filaments arising from the dorsal edge of the foot at the same level as the presetal
lobes. The gill filament is longer than the presetal lobes. The dorsal cirri are
relatively large and arise from the body wall well above the foot in anterior
segments but just at the base of the foot in the posterior part of the body. There
are two long presetal lobes with pointed ends, and the superior one is distinctly
shorter than the inferior. There is only one low rounded postsetal lobe; the
ventral cirrus is short and pointed.
This is a new record for South Africa.
Glycera roux Aud. & M.-E. 1834
Glycera sagittariae McIntosh 1885, p. 346, pl. 42, fig. 8; pl. 22A, fig. 10.
Glycera rouxii. Fauvel 1923, p. 389, fig. 153 a—c.
Glycera goesi Malmgren, McIntosh 1925, p. 69.
Records; TRA.G0(3))727(1); PAL 2973).
Notes: The proboscidial papillae are simple blunt cones some of which
show a trace of two rings. There are also a few spherical papillae. The jaw
supports lack one prong of the complete Y shape. Branchiae start on the 18th
foot as long, single, retractile filaments arising from the anterior face of the
parapodium. There are two, unequal, pointed postsetal lobes, the superior
being slightly longer.
An examination of the type of Glycera sagittariae McIntosh 1885 which is
now in the British Museum shows that it has feet with single retractile gill
filaments arising from the anterior face of the parapodium and that the postsetal
lobes are triangular, the superior one slightly longer than the inferior. Both
postsetal lobes are shorter than the presetal ones. 3
Fauvel (1932) has recorded G. sagittariae but his description and figures do
not agree with the type specimen seen by me. Also MclIntosh’s pl. 42 fig. 8
shows no gill on the dorsal edge of the foot, and his remark that the gill arises
‘from the upper and anterior part of the foot’ agrees with his specimen. Hartman
(1950) suggests that Fauvel’s specimen is really a G. tesselata, but G. tesselata
lacks gills. It differs in the proboscidial papillae from G. alba or G. prashadt.
Hartman suggests that G. sagittariae McIntosh is G. gigantea but this is incorrect,
since the latter has globular gills and rounded postsetal lobes.
Ophioglycera eximia (Ehlers) 1900
Goniada eximia Ehlers, Monro 1936, p. 141, fig. 25 a-j.
Ophioglycera eximia Hartman 1950, p. 38.
Records: AFR.1579(1); FAL.233(1).
Notes: These specimens are identical with Monro’s specimens in the
British Museum. The largest specimen has 4-5 teeth on the macrognaths, 30
dorsal and 19 ventral micrognaths. Parapodia 1-57 are uniramous with a dorsal
cirrus which is notched on the dorsal edge; at the 58th foot the dorsal cirrus is
THE POLYCHAET FAUNA OF SOUTH AFRICA 331
accompanied by a smaller inferior lobe; on the 60th notosetae appear between
_ these two lobes and on succeeding feet the inferior notopodial lobe grows
larger and by the 7oth the two lobes are equal in size.
This is a new record for South Africa.
Glycinde capensis n. sp.
(Fig. 8 f-A)
Records: FB.306(1), 316(2); FAL.209(2), 250(1), 375(1 juv.); MB.81
(1 juv.); SCD.61(1), 63(1), 82(1).
Description: The holotype is the largest of the six incomplete specimens from
False Bay and comes from FB.306. It measures 40 mm. for 112 segments and
is pale yellow in alcohol. The tapered prostomium is 8-ringed with 2 pairs of
eyes and 4 minute terminal antennae. The basal pair of eyes is internal and
the distal pair just below the terminal antennae, is external. The long proboscis
is covered with the usual rows of papillae and is armed with a pair of ventral
_ macrognaths each with 5 teeth and a dorsal ring of 24 micrognaths (one of the
paratypes has only 15).
According to Hartman’s formula the proboscidial papillae (fig. 8f) consist
of 6 groups. Group I which lies along the median dorsal line is formed by a
sparse double row of minute tubercles each with a single point (fig. 8f1).
Group II (fig. 8f) consists of 6 oblique rows of much larger tubercles running
along the dorso-lateral surface. IIa the (dorsalmost) is small and the apex of
each tubercle ends in 2 points. IIb and IIc are the largest and each tubercle
is curved with a single apical point. IId, e and f decrease in size and the apex
of each ends in two points like the open beak of a bird. Group III (fig. 8f?)
consists of a row of minute, oval tubercles each ending in double points. Group
IV (fig. 8f4) is a row of slightly larger tubercles, each with a 3-pointed apex.
Group V (fig. 8/5) is a row of large, soft, bluntly conical papillae each with a
small pore at the apex. Group VI, as usual, is absent.
The anterior region of the body consists of 28 uniramous segments, each
with a straplike dorsal cirrus (fig. 8g), a compressed and tapered setigerous lobe
and a ventral cirrus similar in shape to the dorsal but a little shorter.
All along the length of the body there is a tendency for the parapodial
lobes to become shorter and broader and for the ventral cirrus to be attached
more distally. This is true both for the anterior as well as the posterior region.
Moreover, the rudiment of what, in the posterior region behind segment 29
will become the notopodium becomes evident about the 20th segment as an
inferior thickening and later expansion of the dorsal cirrus. The setigerous
lobe consists of the fused presetal and postsetal lobes with the setae issuing as
superior and inferior fans. — |
_ The posterior, biramous region starts at setiger 29. Here, as the notopodium
develops, the dorsal cirrus becomes shorter and moves into a postero-dorsal
332 ANNALS OF THE SOUTH AFRICAN MUSEUM
position while an anteroventral lobe grows out and soon reaches the same
size. The notosetae issue from a slit between them. The neuropodium is homo-
logous with, and essentially similar to, the setigerous lobe and ventral cirrus of
the anterior region. A typical foot of the posterior region (fig. 8h) has a broadly
triangular notopodium with a small flattened dorsal cirrus and a larger but
still essentially triangular neuropodium with a ventral cirrus.
In the anterior region the setae are all compound with spinigerous blades.
In the posterior region, the notosetae are 2-3 acicular setae with bluntly
hooked ends protected by a spike-like guard. The neurosetae are similar to the
setae of the anterior region.
The South African material is obviously close to G. nordmanni (Malmgren)
and G. wirent Arwidsson. However both of these are northern forms, the type
locality of the former being Norway and of the latter, the Behring Straits.
Moreover there are several small differences from both these species.
G. nordmanni has 36-37 anterior segments and examination of several specimens
of various sizes in the British Museum showed that this figure is surprisingly
constant. G. wireni has 31 anterior segments, only a single pair of eyes at the
base of the prostomium and few (17) micrognaths. It seems safer to distinguish
the South African material as a separate species until more is known about the
variation and distribution of these rather rare worms.
Glycinde kameruniana Augener 1918
(Fig. 8 j-m)
Glycinde kameruniana Augener 1918, p. 398, pl. 4, fig. 93; pl. 7, fig. 211.
Records: FAL.237(1), 341(1), 375(1); SCD.109(4).
Description: As far as I am aware this species has not been recorded since
Augener’s original description of a 10 mm. ovigerous female from the
Cameroons in tropical West Africa. Augener’s description is very incomplete
and his figures add nothing to the text. For this reason a full description of the
South African specimen is given below.
The specimen dredged in False Bay is a complete, ovigerous female
measuring 39 mm. with 100 segments. It is pale in alcohol.
The prostomium has 8 indistinct rings, 4 minute terminal antennae and
a pair of eyes embedded in the basal ring. A distal pair of eyes is lacking. The
proboscis has a pair of ventral macrognaths each with 4 teeth and a dorsal
arc of 4 micrognaths. The papillae on the proboscis (figs. 8j 1-5) are of the
usual type. Those of the mid-dorsal row (group I (fig. 871) are minute oval
tubercles with a single point. Group II (fig. 87?) is a dorsolateral band formed of
5 large falcate tubercles in oblique rows; IIa is small, stout and has a single
point; IId is very large, with a curved single-pointed tip; IIc is similar but the
tip is not so sharp and IId and ¢ are progressively smaller and have two points
curved like the open beak of a bird. Group III (fig. 87%) is a row of minute
THE POLYCHAET FAUNA OF SOUTH AFRICA 333
tubercles each with 3 points. Group IV (fig. 8j*) is a row of slightly larger
tubercles whose oblique tops have 3 points dorsally and a sort of prow
ventrally. Group V (fig. 875) is as usual, a row of large soft papillae with faintly
bilobed apices.
The anterior uniramous region consists of 21 segments. Each parapodium
(fig. 8/7) consists of a strap-like dorsal cirrus, a setigerous lobe with a single
tapering presetal lobe, a similar, subequal postsetal lobe and a ventral cirrus
essentially similar to the dorsal one. The first few parapodia (e.g. the 8th)
have the dorsal cirrus as a simple strap but from about the 15th it becomes
obvious that the inferior side of the cirrus is bulging so that such dorsal cirri
could conceivably be described as having a notch below the tip.
In the posterior region the body is a little flatter and the biramous para-
podia are relatively larger. The notopodium, formed as a ventral outgrowth
from the dorsal cirrus, is at first a simple bilobed structure but further back the
dorsal cirrus becomes relatively shorter and posterodorsal in position. The
setigerous lobe itself develops a minute second lobe (see fig. 8m) The neuropo-
dium remains essentially similar to the setigerous lobe of the anterior region
but here it is evident that the presetal lobe is distinctly longer than the postsetal
lobe.
The setae are of the usual types. In the notopodium there are usually 2
acicular setae (fig. 8k) with bluntly curved apices and pointed guards. The
setae of the anterior region and those of posterior neuropodia are slender-bladed
spinigers. I have not been able to see the detail given by Hartman (1950).
The above description agrees with Augener’s brief account in regard to
the number of anterior segments, setae, shape of parapodia and eyes. Of the
micrognaths he says he was only able to see ‘einige ganz feine schwarze Punktchen’.
... | take this to mean that there were only a few micrognaths. Augener gives
no account of the proboscidial papillae.
_ G. solitaria (Webster) from the Atlantic coast of U.S.A. has been re-
described by Hartman (1950) and is obviously close to the present material.
The number of anterior segments is a little larger (24), a distal pair of eyes is
present and there are more micrognaths (10). On the other hand the papillae
on the proboscis seem generally similar though groups IV and V are a little
different. The parapodia are very alike. It is possible that further collecting on
the tropical West African coast will show that G. kameruniana is a synonym of
G. solitaria.
Goniada maculata Oersted 1843
Goniada maculata. Fauvel 1923, p. 392, fig. 154 a-g. Hartman 1950, p. 20, pl. 1, figs. 7-8.
Records: WCD.5 (1 juv.), 26(1 juv.); TRA.1rro(1 juv.); FAL.240(3),
250(1), 352(1).
Notes: South African specimens are quite typical. The papillae on the
proboscis are as figured by Hartman and the feet agree with Fauvel’s figures.
In juvenile specimens it was noted that while the number of micrognaths
334 ANNALS OF THE SOUTH AFRICAN MUSEUM
remains 3—4 dorsally and 3-4 ventrally, the number of teeth on the macrognaths
may be as low as 4; again the change in the parapodial structure occurs a little
earlier, on the 35th as against the 39—41st foot as usually quoted. Specimens of
all sizes lacked eyes and from the 30th foot on, where there are two fingerlike
presetal lobes to the neuropodium, the superior one is always a little longer than
the inferior one.
Family EUNICIDAE
Subfamily EunicinAE
Eunice vittata (Delle Chiaje) 1828
Eunice vittata. Fauvel 1923, p. 404, fig. 158 h—-n. Day 1953, p. 433.
Records: AFR.691(p), 801(p), 994(p); TRA.56(p), 58(p), 71(a), 152(p);
FB.302(1); FAL.26(1), 29(1), 184(1), 223(2), 243(1), 328(3), 334(7), 338(1),
341(1); MB.9(2), 67(4), 78(1); LIZ.18(4); SCD.22(1), 82(1), 99(3), 100(4),
109(3). |
Eunice floridana (Pourtales) 1869
Eunice floridana. Fauvel 1923, p. 402, fig. 157 a-g.
Records: APR 7G © a7 30) aa ely
Eunice pennata (Miiller) 1776
Eunice antarctica Baird 1869.
Eunice savignyi (non Grube) Ehlers 1g08a, p. 88, pl. 11, figs. 7-13. Hartman 1956, p. 283.
Eunice pennata. Fauvel 1923, p. 400, fig. 156 h-o. Monro 1930, p. 118, fig. 42.
Leodice langi Treadwell 1943, p. 3, figs. 14-18.
Records: AFR.691(1), 707(1); TRA.115(1); FAL.375(2).
Notes: Baird’s type of Eunice antarctica from ‘Antarctic seas’ which is in the
British Museum has been compared with £. pennata from Europe identified by
Fauvel and E. pennata from ‘Tristan de Cunha identified by Monro (1930)
and the present material from South Africa. All are identical. Baird states that
in E. antarctica the gills start on the 8th foot, but examination of the type shows
that there are small single filaments from the third foot. E. pennata has fairly
short tentacles with faintly marked annulations towards the tip. It is close to
E. savignyi Grube from the Philippines but the latter is a tropical shallow
water species with distinctly jointed or even moniliform tentacles. Moreover in
E. savignyi the median tentacle is very long and reaches the 16th setiger whereas
in E. pennata it only reaches the 3rd to 8th foot. Ehlers’s record of E. savignyt
from the Agulhas Bank obviously refers to FE. pennata.
By the courtesy of the U.S. National Museum I was able to examine
Treadwell’s type of Leodice lang: collected by H. Lang from 160 fathoms off
Cape Town. The material, Ref. No. A6o099, consists of a single specimen
from which the jaws have been removed. The gills begin on the grd setiger as
simple filaments, reach a maximum of 8 filaments on the 15th foot and end on
THE POLYCHAET FAUNA OF SOUTH AFRICA 30)
the 37th foot. The setae also agree with those of Eunice pennata. Hartman (1956)
has referred Treadwell’s Leodice langi to E. savignyi following Ehlers’s record as
cited in Fauvel (1932).
Eunice aphroditois (Pallas) 1788
Eunice rousseaui Quatrefages, Fauvel 1923, p. 403, fig. 158 a—g.
Eunice aphroditois Day 1943, p. 432.
Records: LB.157(2), 161(1), 299(c); FAL.51(1), 159(1); LIZ.35(1).
Eunice australis Quatrefages 1865
Eunice australis. Fauvel 1932, p. 139. Day 1953, P- 432-
Records: TRA.135(5), 152(p); FAL.30(1), 302(1), 334(2); MB.49(1),
67(1); LIZ.18(1); SCD.9(1), 32(2), 40(4), 54(3), 58(1), 89(9).
Marphysa sanguinea (Montagu) 1815
Marphysa sanguinea. Fauvel 1923, p. 408, fig. 161 a—h.
Records: LB.299(fc).
Marphysa capensis (Schmarda) 1861
Marphysa capensis. Augener 1918, p. 332, text-fig. 33. .
Records: TB.318(4).
Marphysa depressa (Schmarda) 1861
Marphysa depressa. Day 1953, p. 434, fig. 5 n, p.
Records : LB.299(fc), SCD.63(1).
Marphysa purcellana Willey 1904
Marphysa purcellana Willey 1904, p. 263, pl. 13, fig. 17. Day 1953, p. 435-
Records: FAL.219(1), 223(1); MB.20(1); LIZ.18(1); SCD.58(1).
Notes : This is the second record of this rare species. The head and anterior
segments are reddish brown and the falcigerous setae have long blades.
Marphysa sp.
Records: L1Z.35(1).
Notes: A single specimen of Marphysa which is new to South Africa was
dredged in Algoa Bay (LIZ.35.T). It is 20 mm. long by 0-6 mm. with about 100
segments. The tail end is missing. This small, slender worm may be a juvenile
and for this reason is not described as a new species.
336 ANNALS OF THE SOUTH AFRICAN MUSEUM
The diagnostic characters are as follows: Prostomium bilobed, eyes present,
antennae faintly annulated, a little longer than the prostomium. Gills appear
far back (approximately the 6oth setiger) and never develop more than a single
filament. Setae include superior capillaries and inferior bidentate falcigers in
all feet. Posterior feet have in addition comb-setae with about 12 teeth and
bidentate acicular setae with guards. The acicula are bluntly pointed and all
setae and acicula are pale.
Lysdice natalensis (Kinberg) 1865
Lysidice capensis Grube, McIntosh 1905, p. 40, pl. 3, fig. 13.
Lysidice natalensis Day 1951, p. 40. Day 1953, p. 435.
Records: AFR.967(1); FAL.16(4), 21(p), 44(3), 80(p), 113(p), 122(p),
126(1), 134(1), 156(2), 219(1), 269(1), 371(1); MB.49(1), 56(2), 67(1), 78(1);
LIZ.6(1), 18(1), 27(1); SCD.89(1).
Subfamily ONUPHIDINAE
Onuphis emerita Aud. & M.-E. 1834
Onuphis emerita. Fauvel 1923, p. 415, fig. 163. Monro 1930, p. 128, fig. 47.
Records: TRA.41(3); FB.321(3); FAL.205(2), 211(1), 228(2), 238(7),
242(4), 341(3), 352(2); MB.67(1), 81(1); LIZ.19(1), 24(1), 31(2 juvs.);
SCD.96(1).
Notes: The specimens recorded above vary from juveniles of 25 mm.
(LIZ.31) to well-grown individuals of 75 mm. They agree very well with Fauvel’s
description. Some individuals are pale but others have well-developed brown
pigment pattern identical with that on preserved specimens from Naples now
in the British Museum.
This species was first described from South Africa by Monro (1930) but
his account is slightly inaccurate. I have checked his specimen from False Bay
(British Museum No. 1930.10.8.1792) and find that the ‘little conical tubercle’
which he states is restricted to the first 3 or 4 setigers is actually the setigerous
lobe. The structure which Fauvel 1923, p. 415 and fig. 163. C, calls ‘un petit
tubercle conique entre le mamelon sétigére et la base du cirre dorsal’ is a superior
projection of the presetal lobe. This is well marked on the 5th—1oth foot on a
100 mm. specimen from Naples but is not present on the smaller 23 mm.
specimen described by Monro nor on the present material.
Onuphis (Nothria) holobranchiata Marenzeller 1879
Onuphis (Nothria) holobranchiata. Izuka 1912, p. 106, pl. 11, figs. 10-12.
Records: LAM.11(5), 26(5), 35(2), 39(c), 40(2), 41(2); LB.382(7);
TRA.110(1), 113(1), 135(1), 143(3); FAL.375(5), 376(c), 378(3).
Notes: Most of the specimens were broken but the average size when
complete was probably 4—5 cm. Only faint traces of pigment remain between
THE POLYCHAET FAUNA OF SOUTH AFRICA 337
the parapodia and some worms are completely pale. Eyes are present just
external to the inner lateral occipital antennae. The ceratophore of the median
antenna has 10-12 rings but the other antennae are longer and have cerato-
phores with about 14 rings. The jaws are weakly chitinized and the formula
starting with the main fangs is Mx.I = 1 (left) + 1 (right); Mx. II = (5-7) +
(6-8); Mx. III = (6-7) +0; Mx. IV=7+ 10; Mx. V=1+1. The
tentacular cirri are slightly longer than the peristomial segment.
The dorsal cirrus is cirriform throughout. Gills are present as a single
filament on the first and all succeeding feet to near the posterior end of the
body. As usual the first 5 feet have a cirriform ventral cirrus, and a prominent
setigerous lobe and a cirriform ‘post-setal lobe’. From the 6th foot onwards the
ventral cirrus is represented by a glandular pad, the setigerous lobe becomes
inconspicuous and a presetal swelling appears, though the presetal lobe is
never well marked. The ‘post-setal lobe’ diminishes in size and from about the
20th foot it is an inconspicuous conical projection of the foot partially enclosed
by a dorsal arc of setae. This seems to be what usually happens in the
Onuphidinae.
The first 5 feet bear about 6 pseudocompound setae with bivalve hoods.
These setae are all tridentate but the third tooth may be so minute that unless
it is seen in profile, it may be thought to be absent. There are also 2-3 simple
capillaries. On the 6th foot the pseudocompound setae disappear and the
capillaries develop narrow wings. Two bidentate acicular setae appear in the
10th foot and 2 very fine comb-setae with about 18 teeth are present in the
20th foot although they may be present before this. An average foot from the
middle of the body thus contains 2 pale acicula with very slender tips projecting
from the surface, about a dozen narrow-winged capillaries, 2 fine comb-setae
and 2 bidentate acicular setae.
O. holobranchiata first described from Japan has been recorded from the
Indian ocean by Crossland (1904) and Fauvel (1930, 1932). The pseudo-
compound setae are variously described as bidentate and tridentate. Hartman
(1944) has made a very thorough study of the Onuphidinae of the western
hemisphere and under the name Wothria describes two species NV. elegans
(Johnson) and WN. iridescens (Johnson) which, like O. holobranchiata, also have gills
as simple filaments from the first setiger. She discusses the differences between
the three species on p. 88.
Onuphis (Nothria) geophiliformis (Moore) 1903
Nothria geophiliformis Moore 1903, p. 445, pl. 25, figs. 57-59.
Onuphis geophiliformis Izuka 1912, p. 103, pl. 11, figs. 8-9.
Records: FB.311(10), FAL.219(1), 328(1 juv.).
Notes: All the specimens were broken but the largest was approximately
30 mm. long when complete. Most of the specimens are quite pale in alcohol,
338 ANNALS OF THE SOUTH AFRICAN MUSEUM
but two show brown markings between anterior parapodia and one has the
anterior dorsum uniformly brown. The tube is unknown.
Frontal antennae are ovoid, and the occipital antennae have ceratostyles
3-4 times longer than their ceratophores. The median antenna which reaches
back to the 6th setiger has a ceratophore with 8-9 rings. Eyes are present
external to the inner lateral ceratophores. The tentacular cirri are a little
less than the length of the peristomial segment. The jaws are very pale and soft.
The mandibles have the usual form and the maxillary formula is Mx. I =
(left) 1 + (right) 1; Mx. II = 8 + 9; Mx. III = 8+ 0; Mx. IV. =649;
Mx. V=141.
The parapodia are of the usual (igi Gills as single filaments appear on
the 4th or 5th setiger, and are always longer than the dorsal cirrus from whose
base they arise. They persist over most of the body but are absent from the last
40-50 feet. Each of the first 5 feet has a prominent setigerous lobe but thereafter
the setigerous lobe becomes reduced and is hidden between the presetal swelling
and the cirriform post-setal lobe. The latter also decreases and at the 12th
foot becomes a mere papilla partially encircled by setae. The ventral cirrus is a
tapered cirriform organ for the first 5 feet but thereafter becomes a glandular
pad and merges with the setigerous lobe at about the 15th foot.
The hooded pseudocompound setae of the first few feet are tridentate with
the terminal tooth much longer than the second and third. The winged
capillaries are simple, never compound. Hooded and bidentate acicular setae
appear in the 9th—1oth foot and 2-3 minute comb-setae with about 12 teeth
further back.
The above description agrees well with that of Izuka except that he states
that eyes are absent and comb-setae have 16 teeth. According to Moore (1911)
and Hartman (1944), O. geophiltformis is distinguished from O. pallida Moore
Ig11 mainly by the shape of the pseudocompound setae. In the latter species
the terminal tooth is no larger than the others.
Onuphis (Nothria) conchylega Sars 1835
Onuphis conchylega. Fauvel 1923, p. 415, fig. 164.
Records: TRA.33(1).
Genus DIOPATRA
Although Diopatra has been regarded as one of the most clearly defined
genera of the Onuphidinae, distinguished by the spiral arrangement of the fila-
ments on the branchial trunks and the possession of tentacular cirri, it is shown
below in the discussion of Dzopatra dubia that the separation of Diopatra from
Epidiopatra is by no means simple. Moreover the question as to whether there
is one species of Diopatra or several species is a matter of controversy. Important
discussions will be found in Augener (1918), Fauvel (1933), Hartman (1944)
and Rullier (1958). Augener divided the genus into two main groups of species
THE POLYCHAET FAUNA OF SOUTH AFRICA 339
based on the number of teeth on the comb-setae. Fauvel who gives parallel
lists of species with few or many teeth on the comb-setae from similar regions,
concludes that there is one species with minor variations. Hartman used
various characters to distinguish several species in the western hemisphere.
Rullier agrees with Fauvel. ,
I have examined the South African material reported below and material
in the British Museum which includes 30 samples labelled Dzopatra neapolitana
from various parts of the world and samples labelled with 10 other specific
names. This has shown beyond doubt that there ave several species and that
the synonymy of Diopatra neapolitana in particular is very confused. However,
once the diagnostic characters are recognized, the specific distinctions are
fairly clear. The following characters seem to be of value.
PIGMENT PATTERN
While the intensity of the pigmentation varies from specimen to specimen
and the fainter markings fade with age, the pattern is surprisingly constant and
the more intense diagnostic marks have persisted in spirit in specimens of
D. neapolitana collected in the last century.
PROSTOMIAL APPENDAGES
These are often called tentacles but are here referred to as antennae as
they arise from the prostomium. The frontal antennae are presumable affected
by the method of fixation but in one species at least (D. dubia) they are charac-
teristic, shovel-like expansions while in others they are sausage-like or subulate.
The ceratophores of the five occipital antennae are ringed and the number of
rings seems to have a limited variation within a species e.g. 3-5, 6-8, 9-12,
15-20. In D. dubia and in Epzidiopatra hupferiana the rings develop lateral
projections so that the ceratophores may be said to be branched. Hartman
(1944) has also drawn attention to ‘glandular structures on antennae’ but
unfortunately she does not describe her methods of preparation, and the
picture seen varies with the method used. However, if the clear cuticle is
stripped from the ceratostyle, a number of depressions formed by oval cells
will be found projecting into the thickness of the cuticle. These are usually
arranged in 15-25 longitudinal rows or may be irregularly placed.
JAws
Most workers are agreed that the maxillae are of no diagnostic value and
though this may not be the case, there is variation both in the number of well-
formed teeth and in the interpretation of what are teeth and what are not.
Incidentally there is some confusion in the numbering of the maxillary plates
since the left side has one more plate than the right. Dissection shows that
Mx. IV (which are curved plates) correspond, also Mx. I (the main fangs),
Mx. IT and Mx. V, so that it is Mx. III which is missing from the right side.
The mandibles too are usually disregarded, but in at least one species (D.
monrot described below) the mandibular shafts are characteristically swollen.
340 ANNALS OF THE SOUTH AFRICAN MUSEUM
PARAPODIA
These provide useful characters but the structure of the feet changes over
the length of the body and the real nature of the different parts has not been
realized. Thus in the first 4 feet, a presetal lobe is not developed, but the
setigerous lobe which is oval and compressed has been called a presetal lobe.
Further back in the branchiferous region, a true presetal lobe is formed and
may be symmetrical or asymmetrical with a marked inferior projection.
Towards the end of the branchiferous region the presetal lobe is again reduced
to an insignificant swelling in front of the setae. In the anterior region there is
what has become known as a cirriform postsetal lobe. This later decreases is
size, fuses with the now insignificant setigerous lobe and becomes partially
surrounded by a dorsal arc of setae—it becomes, in fact, a small conical
setigerous lobe but for convenience, the term postsetal lobe will be retained for
anterior feet, and in D. chiliensis it is quite characteristic for there are two
cirriform postsetal lobes instead of the usual one. A somewhat similar arrange-
ment is described by Willey (1905) for Diopatra amboinensis from Ceylon. The
ventral cirrus which is cirriform for the first 4 or 5 feet later becomes a
ventrolateral glandular pad. The dorsal cirrus is always cirriform but diminishes
in size on posterior segments and develops a dorsal branchial trunk on the 4th,
5th or 6th foot. The structure of the gills has been studied by several workers
but it seems that the method of preservation has so much effect that only the
most marked differences remain constant for a species. Thus the number of
branchiferous segments (usually 40-50) may be greater in large specimens and
smaller in juveniles. The largest gill is usually the 3rd—6th but in juveniles it
may be the first. In well-preserved specimens the branchial trunk is itself
spiral but this is not true of compressed specimens removed from a tube. In
most species the filaments have a length equal to 3-4 times the thickness of the
trunk but in one species the basal filaments are hardly longer than the thickness
of the branchial trunk.
SETAE
The pseudocompound setae of the first four feet provide characters of the
greatest importance—they may be unidentate, bidentate or tridentate and the
guards (or hoods) may be well or poorly developed. The winged capillaries of
later feet seem to vary in breadth of wings but the degree of serration at the
base of the wing seems to change along the length of the worm. No significant
departure from the norm has been noted in the structure of the four tapered
acicula which just project from the setigerous lobe. The shape of the bidentate
acicular setae seems to be very constant but the number of teeth on the comb-
setae and the angle at which they are set seems to be diagnostic within limits.
Thus in D. neapolitana there are a few large teeth (4-10), in D. cuprea there are
numerous (15-25) small teeth and in D. musseraensis the blade is rolled up like
a paper trumpet and at certain angles appears to have a stout central tooth as
figured by Augener (1918) and Tebble (1955).
5
THE POLYCHAET FAUNA OF SOUTH AFRICA 34.1
TuBE
In most species the projecting end of the tube is beset by shell fragments
or other foreign objects such as leaves set edgeways on. In some, however, the
tube is composed of hardened mud or sand without shells and Hartman has
described one species with a ringed tube.
As mentioned earlier many records have been referred to the species
‘Diopatra neopolitana which was regarded by Fauvel as being widely distributed
in warmer waters. Among others, most of my own records from South
Africa are incorrect and it is convenient before describing new material to
redescribe a specimen of D. neapolitana from Naples and discuss the tangled
synonymy.
Diopatra neapolitana Delle Chiaje 1825
(Fig. 9 a-g)
Diopatra neapolitana. Claparéde 1868, p. 122, pl. 6, figs. 4 a-h. Ehlers 1868, p. 285, pl. 12, figs.
6-20.
non Diopatra neapolitana Crossland 1903, nec Day 1934, nec Day 1957, nec Tebble 1955, nec Monro
1936, nec Fauvel 1932, nec Wasenberg-Lund 1949.
Material (from Naples) British Museum numbers 1898: 5: 6: 137-9;
1919.11.6.25/26; 1921.5.1. 1873/74; 1876:10:4:41; 1890.6.7.9.13 other samples
in the British Museum labelled D. neapolitana from South Africa, West Africa,
West Indies, various parts of the Indian Ocean and Australia were not this
species.
Diagnosis: Pseudocompound setae with a very small secondary tooth and
well-developed guards; comb-setae with 4-10 teeth; a dark spot in the middle
of the back on branchiferous segments.
Description: The following description is based on three British Museum
specimens 98.5.6.137/9 from Naples. The specimens are very large, measuring
24 cm. by 5 mm. with over 250 segments. The general colour is brown, darker
anteriorly and on the inner sides of the ceratophores of the occipital antennae.
There is a short, dark, transverse, mid-dorsal bar on the anterior margin of
each of the first 10 branchiferous segments (see fig. ge).
The frontal antennae are tapered and fairly long. The ceratophore of
the median occipital antenna has 9-11 rings and is about a third the length of
its ceratostyle which has 20-25 broken longitudinal rows of clear cells projecting
into the cuticle. The tentacular cirrus is three-quarters the length of the median
ceratophore.
The mandibles have calcareous bilobed cutting edges and well chitinized,
straight, dark, tapered shafts. The maxillae are also well chitinized and dark.
The supports are heart-shaped and the main fangs (Mx. I) are strong. The
dental formula is Mx. I (left) = 1 + (right) 1; Mx. IT = 8 + 8; Mx. III =
Be Os ix TV — 7 178; Mx. V — 1 -+ 1. In one large specimen Mx. V =
Seo)
342 ANNALS OF THE SOUTH AFRICAN MUSEUM
The first few feet tend to be bent forward and downward, and the first
three (fig. 9a) are of the usual form described earlier for the genus Diopatra.
The first gill occurs on the 4th or 5th foot. The largest gill is on the 7th or 8th
foot and it extends two-thirds of the way across the dorsum; it has about 10-12
whorls of branchial filaments. Individual filaments of the basal whorls have a
length equal to 3 to 4 times the thickness of the branchial trunk. Succeeding
gills decrease slowly in size to end about the 50th foot. The presetal lobe is
well marked from the 5th to the 2oth foot and in these well-preserved specimens
there is an obvious ventral projection (see fig. 9b).
The first four feet have 2 superior capillary setae and a fan of 5-6 hooded
pseudocompound setae. The latter (fig. 9d) usually possess a very small
secondary tooth but this may be absent. Pseudocompound setae are absent
from the 7th foot and are replaced by winged capillaries which later develop
serrations on the base of the wings. Comb-setae (fig. 9g) appear on the 8th foot
and, tested over the whole length of the body, always have truncate ends with
5-10 coarse teeth. Two bidentate acicular setae (fig. of) appear on the 15th
foot and persist over the rest of the body.
Discussion: Neither of the two descriptions given by Delle Chiaje (1825)
and (1841) are sufficient for more than generic diagnosis, though the notes on
pigmentation vaguely suggest D. neapolitana. Quatrefages (1865) described
D. gallica which according to most authorities is synonymous with D. neapolitana,
but not having seen the type, I must rely only on the published description
which is not sufficiently detailed for D. neapolitana. On the other hand both the
description of Claparéde (1868) and Ehlers (1868) are very good. Claparéde’s
figure 4D of the pseudocompound setae probably represents a broken seta
but 4£ shows an unidentate seta and the text states of the pseudocompound
seta: ‘son extremité se recourbe de maniére a constituer une veritable serpe (4E). Chez
quelques individus cette serpe est bidentée.’ If he had added that the secondary tooth
is usually rudimentary the matter would have been clearer. His figures 4H
and 47 show comb-setae with 7 and 1o teeth.
Ehlers (1868) gives a detailed description of pigmentation and of the mid-
dorsal spot on branchiferous segments. His figures show comb-setae with 8
teeth but the pseudocompound setae were presumably broken off short as often
occurs for he took them to be acicula and does not describe pseudocompound
setae at all. Crossland (1903) who describes a different pigmentation, strongly
bidentate pseudocompound setae and comb-setae with numerous teeth
obviously had a different species in front of him. Fauvel (1923), whose figure
166d shows a comb-setae with few teeth and figure 166f shows a strongly
bidentate pseudocompound seta presumably had more than one species before
him. Fauvel (1930) and (1932) had specimens with numerous teeth to the
comb-setae but the structure of the pseudocompound setae is not stated.
Subsequent authors such as myself Day (1934) and (1957), Monro (1937),
(1938), Tebble (1955) and Wesenberg-Lund (1949) have followed Crossland
and Fauvel.
THE POLYCHAET FAUNA OF SOUTH AFRICA 343
Fic. 9. Diopatra neapolitana: a anterior view of 2nd foot; 5 anterior view of 12th foot; ¢ anterior
view of posterior foot; d, d1 pseudocompound seta of 2nd foot and an unidentate variety;
e 10th segment showing pigmentation; / bidentate acicular seta; g comb-seta.
Diopatra neapolitana var. capensis: h, ht pseudo-compound seta and variation; 7 anterior end
showing pigmentation; j bidentate acicular seta; k comb-seta; / anterior view of 2nd foot.
344 ANNALS OF THE SOUTH AFRICAN MUSEUM
It would appear then that the true.D. neapolitana which occurs at Naples
and presumably elsewhere in the Mediterranean has often been recorded in
error from the Indian Ocean, Australia, South Africa, Pople West Africa
and the West Indies.
NOTE ON SPECIMENS FROM DuRBAN BAy
Record: DBN.26(6).
After the diagnostic characters of the genus Diopatra had been determined,
all earlier collections from South Africa were re-examined, and among others
the material from Morrumbene Estuary, Inhaca Island and Durban Bay
reported by me (Day 1957) as D. neapolitana. The Morrumbene and Inhaca
specimens proved to be D. cuprea but the Durban Bay specimens were D.
neopolitana. The latter had the characteristic pigmentation, comb-setae with
5-9 teeth, the inferior projection from the presetal lobe of the 6th—12th foot
but the pseudocompound setae were more variable than usual. Some were
unidentate, some minutely bidentate and some had a well-developed secondary
tooth.
Diopatra neapolitana var. capensis nov.
(Fig. 9 h-l)
Records: LAM.17(1), 48(4), 55(1), 56(5), 60(3), 63(1); SB.135(1), 181(1),
202(2); AFR.1535(1), 1544(1), Senta TRA.g1(1); FAL.209(1); MB.33(2),
34(1), 37(2), 81(4); LIZ.19(p), 23(3), 24(4); SCD.20(4), 63(10).
Description: The type is one of the 3 specimens from station LIZ.23 dredged
in Algoa Bay at 33°58'S/25°43’E in 38-5 metres on a mud and clay bottom.
It is an anterior half of a worm and measures 50 mm. by 4 mm. for 50 segments.
The tube is of the usual form with broken shells attached edgeways. The differ-
ences between this variety and the stem form concern the colour pattern, the
setae, the shape of the gills and the presetal lobe of branchiferous feet.
The central area of the prostomium between the occipital antennae is
touched with brown and both the inner and outer faces (but not the sides)
of the ceratophores are brown. The peristome (fig. 92) has a continuous
transverse bar and the first five setigers have 5 dorsal marks, 3 on the anterior
margin and 2 posteriorly thus _ _ . Within the next few segments the 3
anterior marks fade but the 2 posterior marks persist to the middle of the
branchiferous region.
The cuticle of the occipital antennae has 25 rather irregular rows of clear
oval cells projecting into it.
The only obvious difference in the shape of the feet between the Cape and
the stem form from Naples, concerns the presetal lobe which, in the stem form,
is well developed between the 6th and 15th feet. In this Cape material the
inferior projection of the presetal lobe is poorly marked or absent so ae te
lobe is symmetrical.
THE POLYCHAET FAUNA OF SOUTH AFRICA 345
The gills appear on the 4th or 5th foot, soon reach a maximum size and
thereafter decrease gradually to end about the 50th foot. Each gill has a
characteristically long stout trunk and short filaments; thus the basal filaments
first appear half-way up the trunk and are not longer than twice the width of
the trunk.
The pseudocompound setae of the first feet (fig. 92) are almost always
unidentate and only on one specimen was a small rudiment of a secondary
tooth found. Moreover this Cape material usually lacks hoods over the apices
of the pseudocompound setae and only in a few cases have vestiges of hoods
been seen. Comb-setae (fig. 9k) with 9-12 rather fine teeth appear about the
12th foot, and bidentate acicular setae (fig. 97) on the 18th foot.
Juvenile and Epidiopatra stages. Five young stages were found which varied
from 15 mm. to 25 mm. in length. Three of them lacked tentacular cirri but
the cirrophores of the occipital antennae were quite normal, 7-ringed and
without lateral branches such as commonly occur in Epzdiopatra species.
Pseudocompound setae were unidentate and provided with small hoods.
No comb-setae had been developed but bidentate acicular setae were found in
posterior feet. Gills were present from the 4th or 5th foot to the gth, roth or r1th.
The first gill was not only larger than succeeding ones but also better developed.
The only markings were brown spots in the intersegmental junctions above
the feet in the middle of the body (cf. D. punctifera Ehlers).
Two other specimens of the same size possessed tentacular cirri and had
better-developed gills from the 4th to the 16th foot. In all other respects they
agreed with the Epidiopatra stages.
This evidence, together with that of Monro 1924A who described a juvenile
Diopatra cuprea from Madeira which also lacked tentacular cirri and the note
below (p. 350) which describes the Epidiopatra stages of Diopatra dubia, shows
that the juveniles of certain species of Diofatra lack tentacular cirri. However,
this is not always the case. Among empty Diopatra tubes dredged from False
Bay was a mud cocoon containing 6 post-larval Diopatras of 8-15 mm. Each
had a normal pair of tentacular cirri, obvious spiral gills and bidentate pseudo-
compound setae.
Diopatra monroi n. sp.
(Fig. 10 a)
Diopatra cuprea (non Bosc) Augener 1918, p. 530, text-fig. 39 (partim).
Diopatra punctifera (non Ehlers) Monro 1930, p. 124, fig. 44 a-b; 1936, p.-147.
Records: SB.124(1), 208(1); LB.162(3); WCOD.26(1); AFR.718(7);
TRA.68(abundant), 70(c), 77(c), 88(8), 89(fc); FAL.352(2).
Diagnosis: Pseudocompound setae strongly bidentate; mandibles stout
and thicker in the middle than at the ends; tube usually of compacted mud.
Description: The holotype is one of 7 anterior fragments from AFR.718.D.
It is 28 mm. long by 4 mm. wide with 66 segments. The complete worm was
346 ANNALS OF THE SOUTH AFRICAN MUSEUM
probably twice this length and not fully grown. The general colour is brownish
anteriorly and pale posteriorly, but the important features are that there is a
dark spot on the prostomium behind the median occipital antenna and the
peristome and anterior segments have brown cross bars across the back which
are best marked in the branchial region (see fig. 10¢).
The frontal antennae are ovoid and the five occipital antennae are borne
on ceratophores with 6-8 rings. The median antenna is 4-8 times the length
of its ceratophore. When the cuticle is skinned off the antenna no clearly
marked rows of cells were found projecting into it. If such cells are present they
must be poorly developed. No eyes are visible on the prostomium.
The mandibles (fig. 105) are quite characteristic. The cutting edges are
normal and white but the supports are black, sausage- or spoon-shaped and
curved. Even in juveniles where the mandibles are pale they are thicker in
the middle than at the ends. The maxillae on the other hand are always
weakly chitinized and brown. The supports are heart-shaped and the main
fangs (Mx. I) are soft. The dental formula of the type is: Mx. I = 1 (left) + 1
(right); Mx. Il=5+ 7; Mx. II]=6+0; Mx. IV=5+7; Mx. V=
I+ 1.
Dissection of other specimens shows that the number of teeth is not
constant so that the average formula is: Mx. 1 = 1 (left) + 1 (right); Mx. II =
(5-7) + (6-8); Mx. III = (6-8) + 0; Mx. IV = (5-10) + (7-10); Mx. V
= 1+ 1. The fifth pair of plates are pale with a single dark tooth on the edge
of each. !
Anterior feet (fig. 10a) each have an ovoid and compressed setigerous lobe,
tapered dorsal and ventral cirri and a tapered post-setal lobe. The presetal
lobe is not developed in the first few feet. The dorsal cirrus persists throughout
the branchiferous region but gradually becomes more slender and reduced in
size. The ventral cirrus is reduced over the first 5 feet and from the 6th onwards,
it is represented by a ventro-lateral glandular pad. The setigerous lobe soon
becomes less obvious and on the 6th foot it is concealed behind a low, swollen
presetal lobe. The postsetal lobe is gradually reduced in size to a conical papilla.
Gills appear on the 5th foot, reach a maximum size by the 8th foot and continue
to the 43rd foot. Gill filaments are of normal length and appear fairly near the
base of the trunk which is not particularly stout.
The first four feet bear a fan of setae including 1-2 superior simple capil-
laries and 4-5 pseudocompound setae (fig. 10e) each with a strongly bidentate
tip covered by a pointed hood. Only the simple capillaries persist and these
become more numerous in posterior segments; the blades remain narrow and
do not develop serrations at the base. From the 6th or 8th foot comb-setae
with 15-25 teeth appear. Bidentate acicular setae with guards (fig. 10/)
appear about the 16th foot.
The tube is of hardened mud or occasionally of sand and no sign of shells
have been seen in many hundreds of specimens examined, even in those from
shallow depths where the substratum is sand and shells. This species occurs on
THE POLYCHAET FAUNA OF SOUTH AFRICA 347
Fic. 10. Diopatra monroi: a anterior view of 2nd foot; 6 ventral view of mandibles; c anterior end;
d comb-seta; e pseudocompound seta from 2nd foot; f bidentate acicular seta.
Diopatra dubia: g anterior view of 2nd foot; A anterior view of 8th foot; 7 anterior end;
Jj anterior view of prostomium with ceratostyles omitted; k pseudocompound seta from 2nd foot.
348 ANNALS OF THE SOUTH AFRICAN MUSEUM
muddy bottoms along the west coasts of South and South West Africa in
enormous numbers. ‘The main banks are 60-100 fathoms deep and fishermen
report that at times the mud tubes may clog a trawl so that the gear has to be
cut away. ,
I have examined the specimens recorded by Monro (1930) and (1936)
as D. punctifera and find that they are identical with the specimens described
here; equally certainly they are not the Diopatra punctifera which Ehlers described
from the Agulhas Bank. Monro himself drew attention to the difference in the
mandibles which are so characteristic.
By the courtesy of the Director of the Hamburg Museum I was able to
examine the three specimens V.8718 recorded by Augener 1918 from Swakop-
mund under the name of D. cuprea. According to Augener they were obtained
from mud tubes, and they still retain a dark spot behind the ceratophore of the
median antenna and brown cross-bars across anterior segments. ‘The mandibles
have stout black spoon-shaped shafts. They are in fact typical examples of
D. monrot.
Diopatra dubia n. sp.
(Fig. 10 g—k)
Records: TRA.73(2), 80(2), 110(1), 113(1), 143(3); FAL.237(3), 240(3),
352(3)s 376(3).
Diagnosis: A small species with poorly developed gills, flattened, shovel-like
frontal antennae and side branches on the ceratophores of the occipital
antennae.
Description: The holotype was selected from a collection of 7 specimens
from FAL.237.H. It is a well-preserved specimen measuring 23 mm. by 1:8 mm.
with 50 segments. The tail end is missing. The tube is rather fragile and com-
posed of flocculent debris, small sand grains and calcareous fragments. The
worm is mainly pale but there are golden brown markings anteriorly though in
other specimens these may be faint or lacking. The type has flecks forming a
rough circle on the slightly concave area of the prostomium between the
antennae and 3 paratypes each have a brown spot in the ceratostyle of the
occipital antennae. The anterior segments are ringed with brown with a break
over the ventral nerve cord and the glandular ventral cirri. The markings are
strongest between the parapodia and persist there after they have faded
elsewhere. The palps are oval cushions projecting outwards below the frontal
antennae.
The frontal antennae (fig. 107) are unusual. Instead of being stout and
cylindrical as in most species, they are flattened, much broader than long and
rather spade-shaped. In large specimens they are fused for most of their
length. |
The occipital antennae (fig. 107) all have 5-ringed ceratophores with side
projections on each ring except the enlarged terminal one. The median antenna
has branches on either side of its ceratophore but the inner laterals and outer’
THE POLYCHAET FAUNA OF SOUTH AFRICA 349
laterals only have branches on their medial sides. The ceratostyles are relatively
short, and the whole median occipital antennae only reaches back to setiger 3.
In fresh specimens an internal brown spot may be seen in the ceratostyle.
The jaws are very weakly chitinized. The mandibles have calcified cutting
edges but their straight, tapered shafts are pale except for narrow black lines
on the inner edge of the dorsal surface. The maxillae are equally soft and only
the teeth, outer margins and a conspicuous crescentric area medial of Mx. IV
is blackened. The supports as usual are heart-shaped and starting with the
main fangs (Mx. I) the formula of the holotype may be expressed as: Mx. I =
(left) 1 + (right) 1; Mx. II1=9-+ 9; Mx. II]=9-+0; Mx. IV=6+09;
Mx. V = 14+ 1. Dissection of other specimens has shown that this formula
is higher than most and the range is shown by the figures in brackets: I = 1
fea) © (right); Il — (5-9) + (6-9); ITT — (6-9) + 0; IV = (5-6) + 9;
Ve r+ 1.
The peristomium bears a pair of rather short but quite obvious tentacular
cirri showing clearly that the type material must be included in the genus
Diopatira as at present defined.
The anterior feet (fig. 1og) are of the usual fom, There is an ovoid
setigerous lobe with a rather short cirriform post-setal lobe behind it. The
cirriform dorsal cirrus above is considerably longer and the cirriform ventral
cirrus below is almost as long.
The dorsal cirrus develops a dorsal gill on the 5th foot. However, the gills
(fig. 10h) are never well developed. The largest gill is the 3rd on the 7th foot
and this has only 2—3 whorls of filaments and just reaches across the mid-dorsal
line. ‘Thereafter the gills diminish rapidly and in most cases there is only a small
dorsolateral tuft of filaments. The last gill occurs on the goth foot. The postsetal
lobe of anterior feet diminishes in size posteriorly, fuses with the setigerous lobe
and disappears about the 25th foot. The cirriform ventral cirrus of the first
3-4 feet becomes a ventrolateral pad on later segments. No presetal lobe was
distinguished.
Apart from the 4 colourless and tapered acicula present in all feet, the
anterior feet each have one superior capillary seta and about 5 pseudocompound
setae (fig. 10k) with long pointed hoods over the usual strongly bidentate tips.
The pseudocompound setae are replaced by winged capillaries on the 6th
foot. These increase to 12 and then decrease again to about 4. Bidentate acicular
setae with guards appear about the 1oth foot and fine comb-setae with 18-20
teeth appear on the 15th.
One specimen (not the holotype) has a complete posterior end which bears
two fine cirri below the anus.
The above description of a small species of Diopatra with shovel-shaped
frontal antennae, branched ceratophores to the occipital antennae and poorly
developed gills would seem at first to be an unusually well-defined species of
Diopatra. However, further samples revealed very similar worms of the same
size which lacked tentacular cirri. Careful study revealed certain small differences
350 ANNALS OF THE SOUTH AFRICAN MUSEUM
detailed below but it was obvious that all these forms were very closely related
if not mere sexual differences or growth forms of the same species. The question
immediately arose as to whether the genera Diopatra and Epidiopatra are distinct.
All the present material was re-examined as well as specimens of Epzdiopatra
hupferiana and E. drewinensis from tropical West Africa and Monro’s material
from False Bay which has been referred by me (Day 1957) to E. hupferiana
var. monrot. It should also be noted that the discovery of a cocoon of Diopatra
neapolitana var. capensis containing newly hatched juveniles 10-20 mm. in
length with well-developed tentacular cirri shows that in one species of Diopatra
at least, these structures are present from the earliest stages. I have finally
decided to leave the matter open for the present and merely give the characters
of the form with tentacular cirri below.
NOTEs ON AN ‘EPIDIOPATRA FORM’ OF DIOPATRA DUBIA
Records: TRA.73(3), 74(1), 80(2).
Notes: This is a form of Duopatra dubia which agrees with the above
description with the following exceptions.
Tentacular cirri are absent. Occipital antennae have ceratostyles which are
2-3 times the length of their ceratostyles. The hoods of the pseudocompound
setae are considerably longer and the gills are even more poorly developed.
These ‘Epidiopatra forms’ differ from Epidiopatra hupferiana var. monrot
in pseudocompound setae, in tube formation, in pigmentation, in number of
gills and in the shape of the frontal antennae which are broad and flat instead
of elongate and cylindrical. Again the ceratophores of the occipital antennae
are much more richly branched. The gills are very similar to E. hupferiana
from tropical West Africa but the other characters are still distinctive and the
tubes are not plastered with shell fragments.
Diopatra cuprea (Bosc) 1802
Diopatra cuprea. Augener 1918, p. 350, text-fig. 39 (fartim). Hartman 1944, p. 54, p. I, figs. 9-14.
Diopatra neapolitana (non Delle Chiaje) Crossland 1903, p. 132, pl. 14, fig. 1. Day 1934, p. 54.
Day 1957, p. 92 (partum).
Notes: I have re-examined my own specimens from Portuguese East
Africa and have compared them with specimens in the British Museum from
the Gold Coast. They all agree with Hartman’s description of D. cuprea
collected on the Atlantic coast of the U.S.A. near Bosc’s type locality (Charles-
ton). The pigmentation is diffuse brown and no clear pattern could be
distinguished, except that there is a dark internal spot at the base of the dorsal
cirri of the first two feet. Tentacular cirri are rather longer than usual, almost
as long as the ceratophores of the long occipital antennae. When the cuticle
was removed from an occipital antenna, 15-20 broken longitudinal rows of
clear cells are seen projecting into it. Pseudocompound setae are strongly
bidentate and have well-developed hoods. Comb-setae have 18-25 teeth.
The presetal lobe of the 5th—15th foot is small and symmetrical.
THE POLYCHAET FAUNA OF SOUTH AFRICA 351
It is very probable that many of the records of D. neapolitana from the
Indian Ocean which refer to specimens with bidentate pseudocompound
setae and comb-setae with numerous teeth really refer to D. cuprea.
Diopatra cuprea var. punctifera Ehlers 1908
Diopatra punctifera Ehlers 1908a, p. 78, pl. 10, figs. 1-11.
Records: ‘TRA.143(2); False Bay: 25 records from 7—64 metres on shelly
sand; MB.81(3); KNY.6(1); LIZ.3(c); SCD.1(2), 25(c), 26(6), 33(1), 50(c),
58(1), 61(c), 63(3), 74(1), 78(2), 80(1), 94(2).
Description: This Cape material differs from tropical specimens of D. cuprea
in several minor respects but these differences are constant and for this reason
it is as well to refer them to a separate variety for the present. The differences
concern the pigment pattern, the cuticle of the occipital antennae and the
comb-setae.
The colour pattern though often faint shows a transverse row of 4 brown
spots across the back of each of the first 4-8 segments near the posterior margin
thus - - - -. In the anterior branchiferous region the two inner spots on either
side of the mid-dorsal line spread and fuse with the outer spots so that the whole
back is brown with a white streak down the middle. When the clear cuticle is
removed from one of the occipital antennae, scattered clear cells are found.
The comb-setae have rather fewer teeth than in the stem form. In three speci-
mens comb-setae appeared on the 6th, 8th and oth foot and in different
parapodia the number of teeth on the comb-setae was 9, 14, 13, 14, 9, 10, 15,
14, 11, 15, 18 giving a range from g—18.
The feet are of the usual form without any distinctive features. In the
first five the setigerous lobe is oblique and in setigers 8-15 the presetal lobe is
symmetrical without any inferior projection.
Ehlers (19084, p. 78) has included D. neapolitana as a synonym of D. cuprea
and records specimens from 16°36’S/11°46’E (off South West Africa),
33°50'S/25°48’E (Port Elizabeth). He states that the colour pattern is variable
and his synonymy suggests that he had more than one species in his
collections.
Diopatra punctifera Ehlers (1908a) recorded from the Agulhas Bank at
35°19'5/20°15’E is not clearly described. Setae of the first 4 feet are stated to be
*hellgelebraun, zusammengesetzt: der dunne Schaft lauft mit einem wenig hakenformig
gekriimmten, 0-06 mm. langen Endglied aus, das mit einfachen gedeckten Zahn endet,
die Deckplatte ist tiber den Endzahn hinaus verldngert ; vereinzelt stehen daneben feine
einfache Capillarborsten (Taf. X, fig. 7)’. Reference to fig. 7 shows neither a simple
capillary nor a unidentate pseudocompound seta but a strongly bidentate
seta. The comb-seta is shown with numerous teeth.
Dr. Hartwich of the Berlin Museum very kindly sent me the type material
of D. punctifera for further examination. There are two specimens, both with
mud tubes without any shell fragments. The colour is faint and although the
352 ANNALS OF THE SOUTH AFRICAN MUSEUM
anterior branchiferous region is brownish, no colour pattern remains nor are
there any ‘eye spots’ between the parapodia to which Ehlers refers.
The median ceratophore has 8 rings and the median occpital antenna is a
little shorter than the inner laterals with a cuticle which shows a few scattered
cells. The tentacular cirri are longer than usual, almost as long as the cerato-
phores. The mandibles have straight tapered shafts and both the mandibles and
the maxillae are weakly chitinized. The maxillary formula is given by Ehlers.
Anterior parapodia are of the usual shape without any peculiarities;
there is no presetal lobe other than a swelling from the 6th setiger onward.
The gills start on the 5th foot, have fairly slender trunks and filaments whose
length is 4-5 times the diameter of the branchial trunk.
The first few feet have 1-2 capillaries and about 4 pseudocompound setae
with well-developed hoods and bidentate tips; the secondary tooth is well
developed but distinctly smaller and more slender than the terminal one.
Bidentate acicular setae of the usual shape first appear on the 15th foot and
comb-setae are present on the same foot. The comb-setae are fairly long and
have 15-18 fine teeth set on the very slightly oblique blade.
Ehlers’s type and my specimens from the same locality agree very well
though the colour pattern of the type has faded and the comb-setae have
rather more teeth than usual.
Epidiopatra gilchristi n. sp.
(Fig. 11 af)
Records: SCD.33(1), 100(10), 103(2).
Diagnosis: A long slender species with a tough, translucent tube which is
sometimes annulated (fig. 11a).
Description: ‘The holotype is an incomplete specimen from SCD.33 Bagi
removed from its annulated tube. It is 55 mm. long by 0-5 mm. wide with more
than 150 segments. There appear to be large eggs at the posterior end but the
tube is so tough that it is impossible to remove them without damage. The body
is creamy brown in alcohol with faint brown spots in the occipital antennae.
The head (fig. 11d) bears a pair of subspherical palps, stout subulate frontal
antennae and five occipital antennae born on 5 ringed ceratophores. ‘The inner
laterals have one or two blunt projections on the basal rings of the ceratophores.
The ceratostyles of the median and inner lateral antennae are three times as
long as their ceratophores and extend back to setiger 5 but those of the outer
laterals are considerably shorter. The head is generally pale in alcohol but
there are brown flecks on the prostomium between the antennae and a row of
ocular specks may be discerned outside the bases of the inner lateral antennae.
The peristome is longer and darker than the succeeding segments. There are
no tentacular cirri.
The jaws are pale and very weakly duaehee The mandibles have white
cutting edges and their shafts are so pale that they are hardly distinguishable
THE POLYCHAET FAUNA OF SOUTH AFRICA 353
from the muscle. They appear to be straight and tapered. The maxillae are
mainly colourless but the teeth are tinged with brown and Mx. V are toothless
crescents of a darker brown. The dental formula is: Mx. I =1 (left) + 1
fee) ix) Ti — 38 -- 7; Mx. Il = 7 + 0; Mx. IV = 6+ ?10; Mx. V =
o + o. The first three feet are ventro-lateral in origin and are directed forwards.
All the lobes are flattened against the side of the body probably due to compres-
sion inside the tube. The first foot (fig. 11e) has a subulate dorsal cirrus, a low
presetal fold, then the setae and then a conical postsetal lobe and an inferior
rounded papilla. The ventral cirrus is short and blunt. The next two or three
feet are generally similar but thereafter the ventral cirrus is reduced to a low
glandular pad and the postsetal lobe decreases in size. The first gill (fig. 11f)
appears on the fourth foot as an outgrowth from the small dorsal cirrus. It has
a stout trunk bearing 6 finger-like filaments in an open spiral. Succeeding gills
are smaller and the last arises from setiger 7 so that there are four pairs of gills
in all. Specimens dredged off Natal (station NAD.10) also had four pairs but
in this case they started on setiger 5. Specimens from SCD.100 and SCD.103
had only one gill on setiger 5.
The post-branchial feet each consist of a fingerlike dorsal cirrus, a rounded
setigerous lobe with a sheaf of setae and minute post-setal papilla and below
this a cushion-like ventral cirrus.
The setae of the first three feet consist of about 8 stout pseudocompound
setae (fig. 115) with strongly bidentate tips and well-marked hoods. The dorsal
cirrus contains 3 fine acicula and the setigerous lobe has 3 stout ones whose
pointed tips just pierce the surface. In the fourth or fifth foot broad-bladed
capillaries appear and by the 6th foot the pseudo-compound setae have gone
and the first comb-seta appears. It has about 12 teeth. Two bidentate acicular
setae appear in the post-branchial feet.
This species is easily distinguished from E. hupferiana v. monroi by the
character of its tube (fig. 11a).
Epidiopatra hupferiana Augener var. monrot Day 1957
Epidiopatra hupferiana Augener var. monrot Day 1957, p. 92.
Records: FAL.219(1), 245(1 juv.), 328(1), 365(1), 376(1), 378(2); MB.
88(1).
Notes: ‘The tubes are fragile and covered with debris,—hydroid stems,
pieces of alga, shell fragments, mud and flocculent matter. The worms are
typical and most have only 3 pairs of gills but a large 60 mm. specimen had 4
pairs. In fresh specimens 4 broad brown streaks extend along the back but
these tend to fade to a uniform brown on preservation in alcohol. I have
examined Monro’s specimen in the British Museum (registered number
1930:10:8:1372) and find it agrees with my type, but the colour has faded and
the ceratophores of the occipital antennae lack lateral branches. The latter
character is variable.
354 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 11. Epidiopatra gilchristi: a Tube; b pseudocompound seta from the first foot; c bidentate
acicular seta; d anterior end; e anterior view of first foot; f anterior view of 4th foot.
Leptoecia antarctica: g pseudocompound seta; h bidentate acicular seta.
Rhamphobrachium capense: i, anterior end; j anterior view of first foot; k, k1 pseudocompound
seta and unidentate variation (LIZ.25); 1 comb-seta.
THE POLYCHAET FAUNA OF SOUTH AFRICA 355
A juvenile with 42 segments measuring 9 mm. (FAL.245.J) was obtained
which probably belongs to this species. The frontal antennae are ovoid, the
ceratophores of the occipital antennae have small lateral branches, tentacular
cirri are absent and no gills have been developed. It is interesting to note
that eye specks are still present behind the inner lateral antennae and the ventral
cirri are cirriform on only the first 3 instead of the first 5 feet as in the adult.
Rhamphobrachium capense n. sp.
(Fig. 11 7—l)
Records: TRA.152(1); FB.307(c), 322(1); FAL.58(p), 117(1), 219(c),
378(1); MB.62(1); ?LIZ.25.T(1); SCD.89(2).
Diagnosis: Gills as single filaments from 30th—4o0th foot; anterior pseudo-
compound setae tri- or bidentate with clawed hoods.
Description: ‘The holotype is a well-preserved specimen with 78 segments
measuring 34 mm. The tail end is missing. It was selected from among 30
specimens dredged in False Bay. The tube is weakly constructed of mucus
with adherent fragments of shells, coralline algae and a few sand grains. Fresh
specimens show dark marks on the prostomium, palps, ceratophores of the
antennae, parapodia and there are two rows of spots on the dorsum. All of
these markings fade in alcohol and the type is colourless apart from tiny black
eyespots at the base of the inner lateral antennae.
The prostomium (fig. 112) is ovoid with cushion-like palps, ovoid frontal
antennae and 5 short stout occipital antennae which just reach back to setiger 1.
Each of these antennae consists of a stout bulbous ceratophore with 2 rings
at the base and a slightly longer subulate ceratostyle. The peristomium is
narrow and the tentacular cirri when laid forward do not reach the bases of
the occipital antennae.
The first few segments are usually tilted upward so that the first two feet
point forward, the 3rd—5th obliquely downward and the rest are normally
lateral. Anterior feet (fig. 117) have subulate dorsal and ventral cirri, a low
presetal lip and two unequal conical postsetal lobes of which the superior is
much the larger. The inferior postsetal lobe disappears after the first 3—4 feet.
Then the superior postsetal lobe is reduced and on the 12th foot it is no more
than a low rounded cone. The dorsal cirrus retains its structure throughout
but becomes reduced in size. Between the 30th and goth segment a single
branchial filament grows out from the dorsal cirrus and soon greatly exceeds
it in length and continues to near the end of the body. The short ventral cirrus
is conical for the first 4 feet, reduced on the 5th and on subsequent segments
it forms a ventral glandular pad.
The first three feet each have a few very small capillaries and 12-18 long
pseudocompound setae (fig. 11k) projecting forward. The apex of each is
bidentate or occasionally tridentate though the third tooth is minute and the
350 ANNALS OF THE SOUTH AFRICAN MUSEUM
bivalve hood which covers the apex terminates in tiny claws. The shaft has two
rows of spinules along the inferior margin. In the single specimen from Algoa
Bay (LIZ.25.T) which is otherwise similar to the Cape material, the pseudo-
compound setae were unidentate or minutely bidentate, the shafts had finer
spinules and the ends of the hoods though bent, were not clawed.
The next few feet (e.g. the 8th) lose the pseudocompound setae and have
about 8 winged capillaries while 2 acicula with long tapered tips project from
the surface. In posterior feet either the capillaries are modified to develop
long slender tips like the acicula, or the acicula themselves become more
numerous and project further from the parapodium. Apart from these very
tapered capillaries or acicula there are 2-3 fine comb-setae (fig. 11/) with
about 12 teeth and 1-2 brown acicular setae with the usual bidentate apex and
guards.
This species differs from those previously described by the position and
nature of the gills and the structure of the pseudocompound setae.
FAlyalinoecia tubicola (Miller) 1788
Ayalinoecia tubicola. Fauvel 1923, p. 421, fig. 166 7-g.
Records: AFR.831(1).
Leptoecia antarctica Monro 1930
(Big ree.)
Leptoecia antarctica Monro 1930, p. 133, fig. 50.
Records: FAL.131(2 juvs.), 159(2); ?SCD.3(8).
Notes : The False Bay material agrees well with Monro’s specimens dredged
off the South Shetland Islands in 1,080 metres. The present specimens are
rather smaller, the only complete individual measuring 23 mm. by 0:8 mm.
for 70 segments. Tubes are missing. The worms are uniformly pale in spirit,
but small eyes are visible external to the bases of the inner lateral occipital
antennae.
The frontal antennae are ovoid to sausage-shaped, the occipital antennae
have ceratophores with 4 rings and the ceratostyles are at least 5 times the
length of their ceratophores. The median antenna which is shorter than the
inner laterals reaches back to setiger 8.
Tentacular cirri are absent and the peristome is about the same length as
succeeding segments.
The first three feet project outwards and downwards, but succeeding
ones change until over most of the body the parapodia are dorsolateral.
Dorsal cirri are always cirriform. Anterior ones are approximately equal to
the segmental length but over the rest of the body they are much shorter and
roughly equal to the setae. The setigerous lobe of the foot is never prominent.
6
THE POLYCHAET FAUNA OF SOUTH AFRICA 357
The postsetal lobe is cirriform for the first 3 feet, then decreases and is not
distinguishable after the 8th foot. A presetal lobe is not developed. The ventral
Cirrus is cirriform on the first three feet and thereafter becomes a glandular
pad which becomes continuous with the setigerous lobe from about the roth foot.
The first 3-4 feet contain about 4—6 pseudocompound setae (fig. 11g)
with bivalve hoods and bidentate apices, the second tooth being much smaller
and more slender than the terminal one. Winged capillaries appear about the
4th foot and by the 8th foot there are 4 capillaries with blades well marked off
from the shafts. Two bidentate acicular setae (fig. 114) appear in the oth foot
and comb-setae with about 14 teeth further back. An average foot in the middle
of the body has 4 winged capillaries, 1-2 fine comb-setae, 2 stout acicula with
fine tapered and projecting tips, and 2 bidentate acicular setae with guards.
The posterior end of the body bears 2 pairs of anal cirri which are a little
shorter than neighbouring segments.
The above description reveals several minior differences from Monro’s
types with which the present specimens have been compared. In particular,
South African specimens have more elongate frontal antennae, they have eyes
and the secondary tooth of the pseudocompound seta is distinctly smaller
and smore slender than the terminal one.
The 8 specimens from station SCD.3 are doubtfully referred to L. antarctica.
The material consists of a number of fine horny tubes up to 40 mm. in length
and 0-5 mm. in diameter attached to a stone. The basal parts of the tubes have
sand grains attached to them but the distal parts are naked and erect. Moreover
several of them are twisted into a fine spiral. The worms inside are of course
more slender than the False Bay specimens but seem to agree in structure.
Monro’s specimens had mud tubes; the tubes of the False Bay specimens
are unknown, and until more is known the identification of the SCD.3 material
is doubtful.
Subfamily LumMBRINERINAE
Lumbrineris albidentata Ehlers 1908
(Fig. 12 a—b)
Lumbrinereis albidentata Ehlers 19082, p. 97, pl. 13, figs. 7-13.
Eeccoras ABN. 736(p); TRA.41(7), 74(2), 80(6), 110(1); 11g(c), 116(1),
143(c), TB.303(1), 309(1); FAL.g5(1), 117(1), 206(1), 228(2), 238(2), 241(1),
243(1), 250(1), 251(10), 328(1 juv.), 345(1), 349(1), 375(1 juv-), 378(1);
SCD.105(1).
Notes: Ehlers’s type was small and incomplete and the present material
which includes numerous specimens of all sizes, now makes it possible to
supplement the original description. The prostomium is conical and there is a
dorsal slit at its junction with the peristomium containing nuchal sense organs.
The jaws are large and in juvenile specimens the first 3-4 segments are expanded
3 58 ANNALS OF THE SOUTH AFRICAN MUSEUM
to accommodate them; in adult specimens this swelling is not noticeable. The
mandibles are heavily calcified and the maxillae are quite characteristic, the
formula being: Mx. I] = 1 + 1; Mx. IT = (2-3) + (2-3); Mx. III = 2+ a;
Mx. IV = 0 + 0; the teeth of Mx. II are often edged with white and in one
juvenile (AFR.736.Q) gave the impression of having a double row of teeth.
Mx. III are very small; Mx. IV are very large plates with a black margin
in which a distinct tooth is not differentiated, but the whole forms a cutting
edge, thus it is represented in the formula as o + o. The maxillary supports
are long and triangular without marked notches at their bases.
Anterior feet (fig. 12a) have lamellate lobes which are longer than deep.
The presetal lobe is at first small, but in middle feet it is considerably larger,
and in posterior feet (fig. 125) it is almost as long as the postsetal lobe. Both
lobes project outwards and upwards but are never as long as the setae, and
much shorter than the posterior lobes of L. bifilaris or L. meteorana. Ehlers’s type
lacked posterior segments so he does not describe this bilabiate condition.
There are long compound hooks from the first setiger changing to simple hooks
at the 30th setiger and persisting to the end of the body. Winged capillaries are
also present from the first foot but these decrease posteriorly so that at the 50th
foot there is only one, but thereafter there are usually one or two in most posterior
segments. The foot contains four yellow acicula.
Lumbrineris cf. meteorana Augener 1931
? Lumbrinereis meteorana Augener 1931, p. 300, fig. 8.
Records: SB.177(1), 199(1); WCD.23(1), 26(2).
Notes: Augener described an anterior and a posterior fragment as follows:
Body very slender. Prostomium conical. Mandibles very pale with a tooth-
shaped process near the symphysis. Maxillary formula: Mx. I=141;
Il=5 +5; W1 = (1 or 2) + (1 or 2); IV = 1+ 1. Third maxillary pla.
with indistinct teeth—possibly 1 or possibly 2. Anterior feet with low presetal
and postsetal lobes. Posterior feet with long threadlike presetal and postsetal
lobes of equal length. Winged capillaries restricted to anterior feet. Hooded
compound hooks in the first 20 feet but replaced by simple hooks over the rest
of the body. Type locality: 17°13'S/11°43’E, off the coast of Angola.
My specimens agree perfectly with the above description except in regard
to the maxillary formula. In my specimens the formula is 1 + 1; 3 + 3;
?2 + Pa;1 + 1. Mx. II have three very stout almost bilobed teeth and Mx. III
is a cutting plate which in some cases shows no teeth at all and in others shows
two small projections. Mx. IV is a relatively large plate with a pale centre and
a dark edge from which a single tooth projects. It may also be added that the
prostomium is oval rather than conical, that the postsetal lobe of anterior feet
is only slightly shorter than the postsetal one, that both lobes increase slowly in
size over the middle of the body but in the last 20 segments or so both lobes
increase enormously to form long threadlike projections. The acicula are pale
THE POLYCHAE1 FAUNA OF SOUTH AFRICA 359
Fic. 12. Lumbrineris albidentata: a anterior view of anterior foot; 56 anterior view of
posterior foot.
Lumbrineris heteropoda var. atlantica: c anterior view of 12th foot; d anterior view
of posterior foot.
Lumbrineris brevicirra: e anterior view of anterior foot: f anterior view of middle
foot; g anterior view of posterior foot.
Lumbrineris magalhaensis: h anterior view of anterior foot; 7 anterior view of middle
foot; 7 anterior view of posterior foot.
360 ANNALS OF THE SOUTH AFRICAN MUSEUM
and the hooks change from compound to simple at the 14th foot which is a
little earlier than stated by Augener.
Since all the other characters agree so well, the difference in Mx. II is
surprising. Augener’s type should be re-examined and it may be that the
anterior end he described does not belong to the posterior region with bilabiate
feet. |
My specimens differ from L. bifilaris Ehlers in the structure of the maxillae
and in having jointed hooks anteriorly. In these characters it is closer to
L. albidentata Ehlers described above but a side-by-side comparison proves that
they are distinct. L. albidentata grows to be a much larger, stouter worm but
even when compared with a juvenile it is evident that L. meteorana is longer
and more slender, that Mx. III are larger plates with less distinct teeth, that
the postsetal lobe of anterior feet is conical not lamellar and that both lobes
of posterior feet are much longer and more threadlike than they are in
L. albidentata.
Lumbrineris heteropoda Marenzeller var. atlantica nov.
(Fig. 12 c-d)
Lumbrinereis heteropoda Monro 1930, p. 137. Monro 1936, p. 154.
Records: LAM.44(1); SB.136(1), 179(1), 183(1), 202(4), 203(10); AFR.
882(1), 1224(1), 1535(1), 1545(1), 1554(1), 1576(1), 1335(1); TRA.68(c),
70(c), 71(fc), 77(c), 80(c); WCD.15(6), 19(3), 21(4), 23(9), 26(45), 28(6).
Notes: These are very large worms, a complete specimen being over 300
mm. long, 5 mm. wide and iridescent reddish brown in colour when alive.
The prostomium is short and conical and the maxillary formula as usual
is Mx. T=1+1; Mx. I1=4-+5; Mx. il=2-+2; Mx. IV =f
but the secondary tooth on Mx. III is poorly developed. The maxillary supports
are heart-shaped and nearly as broad as long.
In anterior feet (fig. 12c) the presetal lobe is short and swollen while the
postsetal lobe is ear-shaped and as deep as long. The feet soon increase in
length and the postsetal lobe becomes relatively longer until at about the 6oth
foot it reaches the tips of the capillaries as a finger-shaped organ. In the posterior
part of the body (fig. 12d) it is much longer than the setae and often three times
as long as the basal part of the parapodium.
In anterior feet the setae are all winged capillaries with brown acicula
embedded in the flesh. The superior group of capillaries have very long slender
blades. Short-bladed simple hooks appear about the 4oth foot and for the next
30 segments, both capillaries and hooks are numerous. Thereafter both types
of setae become less numerous and in posterior feet there are about 4 hooks
and one capillary. It is stressed that capillaries may be found even near the
end of the body. The present material has been checked as identical with
specimens described by Monro (1930) from Tristan da Cunha and, as has been
THE POLYCHAET FAUNA OF SOUTH AFRICA 361
shown earlier (Day 1957, p. 94) there are small but constant differences from
material recorded in the intertidal zone of the tropical Indian Ocean.
Lumbrineris cavifrons Grube 1869
Lumbrinereis cavifrons. Day 1953, Pp. 437, text-fig. 6 a—d.
Records: TRA.152(1); False Bay: 30 records from o-35 metres on rock,
gravel and shelly sand; MB.49(5), 57(1), 85(1), 87(4); LIZ.2(1), 18(3),
29(2), 35(2); SCD.40(1), 89(1).
Lumbrinerts latreilli Aud. & M.-E. 1833
Lumbrinereis latreilli. Fauvel 1923, p. 431, fig. 171 m-r.
Records: FB.307(1), 319(1); FAL.334(1); TRA.132(5); SCD.50(1).
Lumbrinerts tetraura (Schmarda) 1861
Lumbrinereis impatiens Claparéde, Fauvel 1923, p. 429, fig. 171 a-i.
Lumbrinereis tetraura Day 1953, P- 435-
Records: LAM.22(1), 35(10), 38(1), 49(1), 52(1); SB.189(10) ; LB.300(c);
SH.204(1), 415(1); TB.320(1); WCD.21(1); FB.331(1); FAL.58(p); LIZ.2(6),
=7(1)-
Lumbrinerts coccinea (Renieri) 1804
Lumbrinereis coccinea. Fauvel 1923, p. 432, fig. 172 g—n. Day 1953, p. 436 with synonymy.
Records: TRA.122(1); WCD.8(1); FAL.8(p), 113(2), 126(3), 127(1),
144(1), 156(7), 162(3), 171(4), 182(1), 214(2), 269(2), 275(4), 327(1), 371(1).
Lumbrineris harimani Day 1953
Lumbrinereis harimani Day 1953, p- 437; fig. 6 e—m.
Records: FAL.245(1); MB.23(1), 88(1); LIZ.19(7), 25(3); SCD.58(4),
89(1).
Notes: Some of these specimens are much smaller than the holotype and
the simple hooks appear as early as the 23rd segment as against the 45th in
the type.
Lumbrineris brevicirra (Schmarda) 1861
(Fig. 12 e-g)
Notocirrus brevicirrus Schmarda 1861, p. 117.
Lumbriconereis brevicirra Ehlers 1904, p. 35, pl. 4, figs. 13-20; pl. 5, figs. 1-2.
Records: TRA.73(1); FAL.359(1).
Notes: The present specimens lack a posterior end. The prostomium is
short and conical with a nuchal pocket at the junction with the peristome. The
362 ANNALS OF THE SOUTH AFRICAN MUSEUM
maxillary formula is 1 + 135 + 5; ?2 + ?2;1 + 1. Mx. III are cutting plates
with 1-2 indistinct teeth. ‘The maxillary supports are heart-shaped.
Anterior parapodia (fig. 12e) are small and each has a poorly developed
presetal lobe and a well-developed postsetal lobe which is compressed, deeper
than long and roughly triangular with a superior point, rather like a dog’s ear.
Towards the middle of the body (fig. 12f) the whole foot grows longer, the
presetal lobe becomes obvious and the post-setal lobe is reduced; further back
still it is similar to, and not much longer than the small pointed presetal lobe
(fig. 12g). The tail end of the worm is missing.
The anterior setae include both simple hooks and winged capillaries. The
capillaries which have very long slender blades, start on the first foot and
continue to the middle of the body (about segment 50). The simple hooks
appear about the 12th foot and continue to the posterior end (segment 120).
The blade is at first very long so that the anterior hooks give the impression of
being capillaries with broken tips, but further back the blade decreases in
length until it is only 2-3 times as long as broad. The acicula are pale
throughout.
This South African specimen has been compared with a New Zealand
specimen in the British Museum (No. 1928.2.29.156) identified by Benham
(1927). Unfortunately the anterior setae are broken. Ehlers (1904, p. 36)
states that Mx. II have 5 teeth on the left and 7 on the right, an unusually
high number. However, his figure (p. 5, fig. 1) suggests that the number is
smaller as do Schmarda’s original drawings.
Lumbrineris magalhaensis Kinberg 1864
(Fig. 12 h-7)
Lumbrinereis pettigrewi McIntosh 1885, p. 239, pl. 36 figs. 7-9; pl. 17A, figs. 11--15, text-figs. 4-6.
Lumbrinereis magalhaensis. Ehlers 1897, p. 74. Monro 1930, p. 135. Hartman 1948, p. 93, pl. 14,
figs. 1-3.
Records: FAL.352(1). McIntosh’s record of L. pettigrewt is station 141
dredged in 98 fathoms off the Cape at 34°41’S/18°36’E.
Notes: McIntosh’s type of L. pettigrewi is in the British Museum. An
examination showed that the prostomium is long and conical; the dental
formula is 1 + 1;4 + 4;1-+ 1; 1-+ 1. The maxillary supports are short and
broad with practically no notch at the base, Mx. III are curved cutting plates
without a distinct tooth, and are best represented in the formula as I + I.
The presetal lip of anterior feet (fig. 12h) is a low ridge. The postsetal lobe of
anterior feet is flattened, has a rounded edge and is deeper than long, but
further back (fig. 122) it becomes more regularly digitiform. Even in the
posterior part of the body (fig. 127) it is much shorter than the setae. The
presetal lobe increases in size but remains a little shorter than the postsetal
lobe throughout. The majority of the setae are broken, but one or two com-
THE POLYCHAET FAUNA OF SOUTH AFRICA 363
pound hooks remain in the 12th foot of one specimen and a few simple hooks
without swollen ends were found in the posterior feet of another specimen.
The acicula are yellow. According to McIntosh the capillaries which are very
long and slender are restricted to the anterior part of the body. However, his
account is very confused for he figures simple hooks in anterior foot of ‘a variety’,
black acicula in one specimen and pale ones in another. Monro 1930 recorded
L. magalhaensis from South Georgia and his specimens which were examined
in the British Museum were found to be practically identical with McIntosh’s
L. pettigrewt. However, the following minor differences were noted. The
prostomium is conical but short, the dental formula is the same, but the
maxillary supports are slightly longer being 1-5 times as long as broad. In these
complete specimens it may be seen that compound hooks are present from the
first setiger. The postsetal lobe of anterior feet is again very deep but here not
deeper than long. Specimen FAL.352 is an anterior half of a worm which has
been compared with the type of L. pettigrewt and appears to be identical. In
this fresh specimen the setae are unbroken and it can be seen that the compound
hooks start in the first foot and persist to the 19th where they are replaced by
simple hooks. All hooks have short blades. The parapodial lobes are short
throughout and in posterior segments the presetal lobe is only slightly shorter
than the postsetal one.
Ehlers (1897) described two forms of prostomium, one long and one short
but both conical. This type of variation, probably due to the method of
preservation, is quite common in the genus. Ehlers however has made one
error in his description. He states that Mx. IV has two teeth. Both McIntosh’s
specimen of L. petiigrewi from the Cape and Monro’s specimen of L. magalhaensis
from South Georgia have Mx. IV in the form of a cutting plate with an
undulating edge, but not two distinct teeth.
_ Hartman (1948) has redescribed Kinberg’s type material of L. magalhaensis
which consists of several specimens. One was without jaws and obviously
dissected by Kinberg. In other characters however, this specimen agrees with
the description given above, and Kinberg stated that L. magalhaensis has
Mx. III with one tooth, and this is the interpretation accepted by Ehlers and
Monro. Hartman dissected other specimens of the type material and found
that in these Mx. III has 2 teeth and the maxillary supports are twice as long
as broad. I suggest that these specimens are different from the one dissected
and described by Kinberg and seem closer to L. latreilli.
Arabella tricolor var. caerulea (Schmarda) 1861
Arabella iricolor var. caerulea. McIntosh 1904, p. 46, pl. 4, figs. 16-17. Day 1953, p. 439, fig. 6n.
ihecords:, ¥B.305(1), . 319(3); FAL44(1),. 51(1), 58(2), 69(1), 80(2),
235(1), 245(1), 249(2); MB.23(1), 41(1), 42(1), 49(2), 56(5), 67(1), 85(1);
LIZ.29(1); SCD.40(2), 89(6).
364 ANNALS OF THE SOUTH AFRICAN MUSEUM
Arabella mutans (Chamberlin) 1919
Arabella mutans. Monro 1933, p. 501.
Records: FAL.184(1), 229(p).
Notes: These worms have slender bodies with the segments as long as
broad. The prostomium is large and oval with four eyes in a line just in front
of the prostomium/peristomium junction. The mandibles are strong and black;
the maxillae have long filiform supports and a dagger-like median appendage.
The first pair of maxillae do not form strong hooks, but all maxillary plates
have the anterior tooth stronger than the succeeding ones, the formula being:
Mx. 1=8+ 8; Mx. T=7+ 7; Mx. TIl=6+6; Mx) 1Vi22e
Mx. V=1-+1.
Parapodia have a pimple-like dorsal cirrus and no ventral cirrus. The
foot has a low rounded presetal lobe and a fingerlike postsetal lobe. Between
these are 3-4 winged capillaries with serrations at the base of the blade and
two acicula with projecting filiform tips.
Drilonereis falcata Moore 1911
(Fig. 13 a-e)
Drilonereis falcata Moore 1911,.p. 298, pl. 20, figs. 150-154. Hartman 1944, p. 179.
Drilonereis filum (non Clap.) Monro 1936, p. 158.
Records: FAL.219(1), 352(1).
Notes: A single anterior fragment of 66 segments was obtained. The
prostomium (fig. 13a) is depressed, oval in plan and lacks external eyes though
there is a faint suggestion that internal eyes may be present. The mandibles
(fig. 13c) are stout, black and roughly triangular with a short hinge line. The
maxillae (fig. 13b) have long filiform supports which are very narrow where
they join the main fangs (Mx. I) and a dagger-shaped median piece which is
blackened throughout. Mx. I are stout hooks with toothed bases, Mx. II are
rectangular with the first tooth rather larger than the rest, Mx. III have an
anterior large fang-like tooth and small denticles behind, Mx. IV and V which
are very close together, each consist of a single fang. In the following dental
formula the difference in size of teeth is not indicated as is sometimes done.
Mx. I=8+6; T=8+ 6; Wl=4 +3; IV =1+4 1; VV —= ee
The first of the two achaetous segments is largely fused with the prostomium
but leaves a crescentric depression on the dorsal surface. Anterior feet (fig. 13d)
are small and may be partially retracted but succeeding ones rapidly increase
in size. There is no dorsal cirrus. The presetal lobe of the foot is a low semi-
circular ridge at the base of the bluntly conical postsetal lobe. From the groove
between the lobes project a fan of about 6 winged capillaries and the filiform
tips of fine acicula may just be seen. A stout yellow aciculum appears on the
18th—24th foot. The parapodia elongate posteriorly (fig. 13e) and the aciculum
THE POLYCHAET FAUNA OF SOUTH AFRICA 365
(sometimes 2) projects further until it almost reaches the end of the conical
and rather short postsetal lobe.
The above description agrees with Moore’s figures and description apart
from minor details. There are more teeth on the main fangs though Moore
mentions some obscure crenulations ‘as well as 3-4 distinct small teeth’.
Hartman states that there are numerous teeth at the base of the forceps.
Again Moore shows one large tooth only on Mx. III whereas here there are
2-3 denticles as well, but Hartman states that Mx. III have 4-5 smaller teeth
as well as the longer one.
Monro (1936) described a specimen dredged off the Falkland Islands which
he referred to D. filum while noting the differences from typical European
forms of that species. I have examined the specimen in the British Museum,
whose registered number is 1936:2:8:2355, and find it to be D. falcata so that
the range of this species is now from California (o—-172 fathoms) to 46°18’S/
65°02’W (100 m.) and False Bay (18-88 m.).
Drilonereis monroi n. sp.
(Fig. 13 f-2)
Drilonereis sp. Monro 1930, p. 142.
Records: AFR.718(1), 801(1), 1319(1), 1535(1), 1544(1), 1545(1), 1554(1),
1576(1), 1581(1); TRA.68(2), 70(1), 77(2), 88(1), 89(1); FAL.237(1), 240(1).
Description: ‘The type was obtained from station AFR.718. It is 220 mm.
long by 3 mm. broad for 250 segments. It is rusty red in colour and extremely
tough and wiry. The prostomium (fig. 13f) is depressed, broadly triangular
and lacks eyes. The pharynx usually protrudes slightly. The mandibles are
lacking, there being merely toughened skin on the floor of the mouth. The
maxillae (fig. 13g) have long filiform supports and an unusually short, shield-
shaped median piece. Mx. I are stout hooks not denticulate at the base and
Mx. V are missing, the formula being Mx. I =1-+ 1; Mx. II = (6-8) +
(6-8); Mx. III = 3 + 3; Mx. IV = 1 + 1. On Mx. II and III the first tooth
is much larger than the others.
Anterior parapodia (fig. 13h) are small but the feet increase in size
posteriorly. There are no dorsal or ventral cirri. The presetal lobe of the foot is
a low curved ridge and the posterior lobe is a short blunt cone. Between these
project a sheaf of winged capillaries and a stout, blunt, yellow aciculum which
is evident even on the first foot. Posterior feet (fig. 137) are longer, the superior
part of the presetal lobe being more expanded, and about half as long as the
postsetal lobe.
The Drilonereis sp. described by Monro 1930 from Tristan da Cunha has
been compared with these South African specimens and is identical.
This species is related to D. nuda Moore described by Hartman (1944)
from California and D. major Crossland (1924) from Suez and Zanzibar, both
366 ANNALS OF THE SOUTH AFRICAN MUSEUM
Fic. 13. Drilonereis falcata: a anterior end; b maxillae; c mandibles; d anterior foot;
é posterior foot.
Drilonereis monroi: f anterior end; g maxillae; A anterior foot; i posterior foot.
THE POLYCHAET FAUNA OF SOUTH AFRICA 367
of which lack distinct mandibles. It differs from D. nuda in not having teeth at
the base of the pincers (Mx. I) and in these respects it is closer to the type of
D. major which I have dissected and whose dental formula is Mx. I = 1+ 1
(main fangs only); Mx. II = 6 + 6; Mx. III = (3-4) + (3-4); Mx. IV =
3 + 3. However, in D. major the teeth on Mx. II are of fairly even size while in
D. monroi the first tooth is much larger than the rest. There also tend to be
fewer teeth on Mx. III + IV but these plates are more variable in all species.
Other differences are in the setae which are always longer in D. monroi.
Moreover the projecting aciculum of D. monroi appears in the first foot and in
D. major in the 15th. It may also be noted that in the posterior feet of D. major
the presetal lobe remains rudimentary while in D. monrot it becomes enlarged, —
nearly as long as the postsetal lobe.
Notocirrus australis n. sp.
(Fig. 14 a—d)
Records: FB.306(1); FAL.22090(1).
Description: ‘The type material from False Bay includes five fragments,
two anterior and three posterior ends. It is estimated that a complete worm
would measure about 100 mm. by 1 mm. with about 200 segments each about
three times as broad as long. The colour is uniformly pale in alcohol.
The prostomium (fig. 14c) is conical with 4 eyes in a transverse row at the
posterior margin. The outer pair is larger than the inner pair. ‘The jaws consist
of well-developed mandibles and maxillae. Each mandible (fig. 14a) is strong
and triangular and the two are narrowly joined in the median line. ‘The maxillae
(fig. 14b) consist of five pairs of toothed plates so closely crowded together that
it is difficult to distinguish the teeth of one plate from those of the next; in
fact plates I and II overlie one another in one specimen and in the other,
plates I and II are fused on the left side. The anterior tooth of each plate is
hooked and slightly larger than succeeding ones but not to the extent seen in
the genus Drilonereis. The supports (or ‘carriers’) are long and slender and the
median piece is very faint and pale. ‘The maxillary formula in one specimen is
Mae F — 7 (left) + 7 (right); I= 7+ 8; WW =7+6;1IV—=5+4; V=
1 + 1 and in the other where Mx. I and II are fused on the left side: Mx. I
ee — i: (left) + 7-and 9 (right); II] = 9 + 6; IV = 6 + 4;
ee ales ae
The first two apodous segments are rather shorter than the subsequent ones,
all of which bear well-developed parapodia of increasing size. Each parapodium
(fig. 14d) has a minute, pimple-like dorsal cirrus above the setigerous lobe. The
presetal lobe is rudimentary and the thumb-shaped postsetal lobe is at first
almost ventral in position, but in posterior segments it moves round to the
normal posterior position. Each parapodium bears about 3 winged capillaries
368 ‘ANNALS OF THE SOUTH AFRICAN MUSEUM
and a stout yellow needle-like aciculum which projects almost as far as the
postsetal lobe. The capillaries have rather broad wings which are smooth except
for a few serrations at the base.
The genus Wotocirrus has most recently been reviewed by Hartman (1944).
The present species is closest to NV. lorum Ehlers from the Magellan area and
N. californiensis Hartman from Southern California. By the courtesy of the
director of the Hamburg Museum I have been able to examine Ehlers’s type.
Unfortunately the jaws have been removed and I have nothing that I can add
to the original description. The main difference between the three species lies
in the dental formula of the maxillae. As has been mentioned, these plates are
small, crowded together and overlapping, and thus difficult to read. NV. lorum
is reported to have only 4 pairs of maxillary plates but Ehlers’s figure 125
suggests that the 4th and 5th dental plates have not been separated. Again it
may be that the left MX. II on which three teeth are shown is partially covered
by (or possibly fused to) Mx. I. WV. californiensis is very close to the present species
except that Mx. II has 13 teeth on the left side and the usual minute dorsal
cirrus is not figured above the parapodium. It is probable that further work will
reveal that the maxillae are more variable than has been suspected and a
number of species will be sunk in the synonymy.
Fauvel (1923, p. 451) regarded WV. scoticus McIntosh 1869 as a doubtful
species but a re-examination of the type material now in the British Museum
(registered numbers 1921-5-1-1681-86) shows that it is definitely a WNotocirrus
though MclIntosh’s description of the structure of the feet is very confused due
to his having inverted his preparation. ‘Thus what he described as a dorsal
cirrus is really the postsetal lobe, and what he referred to as a ventral cirrus is
really the dorsal cirrus. According to Ehlers (1875, p. 55), V. scoticus isa synonym
of N. tricolor (Johnston) 1865. A brief summary of the characters of the type of
WN. scoticus may now be given. Body brown, rather small for the genus and
segments markedly moniliform for the segments are about as broad as long
with deep intersegmental constrictions between one and the next. Prostomium
conical with two pairs of eyes which fade in alcohol. Jaws consist of a pair of
well-developed, triangular mandibles and 4-5 maxillary plates with the usual
long supports. The dental formula is doubtful. McIntosh (1910) gives a
drawing (p. 62, fig. ga) which shows a number of larger and smaller teeth
which may be variously interpreted. One interpretation is Mx. I = (left) 7
+ (right) 8; Mx. II = 12+ 7; Mx. III] =6-+ 7; Mx. IV = 5 + absent;
Mx. V = 1 + absent. The dental apparatus on which his drawing was based
has not been preserved; indeed all the jaws of the type material are missing
except one from the Porcupine Expedition (registered number 1921-5-1-1685).
‘These jaws are very small and, as usual, difficult to read. My reading is Mx. I
= (left) 7 + (right) 8; Mx. I] = 6 + 7; Mx. WI = 5 + 5; Mx. IV =3 +4;
Mx. V = doubtful. It will be obvious that the dental formula quoted depends
on the inclusion or omission of minute or partially formed denticles on the
maxillary plates quite apart from individual variation. The distinction between
THE POLYCHAET FAUNA OF SOUTH AFRICA 369
oe
Fic. 14. Notocirrus australis: a mandibles; b maxillae; c anterior end; d middle foot.
Drilognathus capensis: e mandibles; f vestiges of maxillae; g anterior end; A middle
foot; 7 posterior end.
370 ANNALS OF THE SOUTH AFRICAN MUSEUM
individual species of the genus Wotocirrus rests largely on the dental formula
so that one becomes sceptical as to whether there really are several species and
not one world-wide one.
Drilognathus capensis n.g. et sp.
(Fig. 14 e-1)
Records: 5 specimens found in the body cavity of Onuphis holobranchiata
dredged in Lamberts Bay 18.1.57 (station LAM.11).
The holotype is a complete specimen which is twisted, but if straight would
measure about 3 mm. by 0-3 mm. with about 60 segments. The whole worm
is creamy white, and is tapered posteriorly.
The prostomium (fig. 14g) is well marked off from the succeeding segment.
It is ovoid, somewhat tapered anteriorly and lacks appendages. No eyes are
visible on the surface but when cleared in glycerine two eyes are visible
posteriorly. Dissection of one specimen showed that the mandibles (fig. 14¢)
are well developed and black. They are of the usual Drilonereis type and there
is no sign of the recurved rostra which has been described for Labrorostratus
parasiticus. The maxillae (fig. 14f) are represented by a blacked cuticular
ridge on the dorsal wall of the pharynx. The length of this black cuticular
streak is reminiscent of the long maxillary supports of Drilonereis and Arabella
but no fangs or distinct maxillary plates were seen.
The first two segments lack parapodia. Each of these achaetous segments
is about 4 times as broad as long. Succeeding segments up to the middleof the
body have well-developed parapodia (fig. 14h) of the usual Lumbrinereis type,
though the presetal lobe is rudimentary and even the postsetal lobe is no
more than a blunt cone. From the middle of the body onwards the parapodia
are progressively reduced, first to mere lateral papillae and eventually over the
last 10-15 segments they are entirely lacking. The pygidium however is well
developed (fig. 147) and has a pair of large ventro-lateral lobes projecting
outwards at right angles to the body.
Each parapodium is supported by a stout aciculum of the Drilonereis type.
It is a bluntly pointed yellow needle which in most parapodia seems not to
pierce the skin, but in some it does, and then just projects in front of the post-
setal lobe. There are no other setae, a fact which immediately distinguishes this
species from related genera.
Pettibone (1957) gives a most useful key to endoparasitic members of the
Arabellidae. It is with considerable hesitation that I name this as a new genus,
for according to Pettibone the young stages of Notocirrus occur as parasites in
the Onuphidae and Pettibone’s figures 5L and M of the jaws of Notocerrus ?
spiniferus are not unlike those of the present specimens. However, all the other
genera that have been described have setae in the parapodia whereas
Drilognathus has merely a well-formed aciculum.
THE POLYCHAET FAUNA OF SOUTH AFRICA 371
Subfamily DorvILLEINAE
Dorvillea neglecta (Fauvel) 1923
Staurocephalus neglectus Fauvel 1923, p. 447, fig. 179 1-g.
Dorvillea neglecta Day 1953; p. 439-
Records: SB.183(1); SH.366(1).
Dorvillea egena (Ehlers) 1913
Stauronereis egena Ehlers 1913, p. 501, pl. 35, figs. 1-6. Augener 1918, p. 377, pl. 5, fig. 102,
103, text-fig. 40.
Records: FAL.284(1).
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Monro, C. C. A. 1933. Notes on a collection of Polychaeta from South Africa. Ann. Mag.
nat. Hist. (10) 11, 487-509.
Monro, C. C. A. 1936. Polychaete worms. II. ‘Discovery’ Rep. 12, 59-198.
Monro, C. C. A. 1937. Polychaeta. Sci. Rep. Murray Exped. 4, 243-321.
Monro, C. C. A. 1938. On a small collection of Polychaeta from Swan River, Western Australia.
Ann. Mag. nat. Hist. (11) 2, 614-624.
Moore, J. P. 1903. Polychaeta from the coastal slope of Japan and from Kamchatka and
Bering Sea. Proc. Acad. nat. Sci. Philad. 55, 401-490.
Moore, J. P. 1911. The polychaetous annelids dredged by the U.S.A. Albatross off the coast of
southern California in 1904. III. Euphrosynidae to Goniadidae. Proc. Acad. nat. Sct.
Philad. 63, 234-318.
Moreans, J. F. C. 1959. The benthic ecology of False Bay. Part I: The biology of infratidal
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387-442.
PetTiponE, M. H. 1948. Two new species of polychaete worms of the family Polynoidae from
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PeTTIBONE, M. H. 1957. Endoparasitic annelids of the family Arabellidae with descriptions of
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Ramsay, L. N. G. 1914. Polychaeta of the family Nereidae collected by the Scottish National
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WILLEy, A. 1904. Littoral Polychaeta from the Cape of Good Hope. Trans. Linn. Soc. Lond.
(2), 9; 255-268.
WILLey, A. 1905. Report on the Polychaeta collected by Professor Herdman at Ceylon in 1902.
In Herdman, W. A. Report to the government of Ceylon on the pearl oyster fisheries . . . 4, Suppl.
Rep. 30, 243-324. London: Royal Society.
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ANNALS
_ SOUTH AFRICAN MUSEUM
PART IV, containing :—
Modern Giraffes and the Fossil Giraffids of Africa. By RONALD SincEr, and
EpouarpD L. Bone. (With Plates I to LII and 27 figures in the text.)
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MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA?
RONALD SINGER?
Anatomy Department, University of Cape Town?
and
Epouarp L. Bonrt
Palaeontology Department, Geological Institute, University of Louvain, Belgium
The publication of this paper was in part made possible by a grant from the
‘Fondation Universitaire’ of Belgium and the South African Council for
Scientific and Industrial Research.
CONTENTS
PAGE PAGE
GENERAL INTRODUCTION . : , ee TAs Olduvai (Oldoway) Gorge, Tanganyika. 415
Sees Gavinc CIRARSE 376 Orange Free State (Union of South Africa) 445
Pe toction ; : 6 Makapansgat (Northern te Union
; : PE: of South Africa) ; : x AT
General description of eiratid dentition 376 H ‘
opefield (Cape Paine, Union of
List of recent Giraffa camelopardalis studied 380 South Africa) . } . 472
Observations on tooth eruption and
cranial sutures E j . 382 GENERAL Discussion AND CONCLUSIONS . 49QI
Variations of the cusps of ue tooth crown 390 Summary of observations on and
Dimensions of the teeth : E . 304. variations in Sivatherines . . 492
Baaeanemalies ; er 4Od: Diagnosis of African Fossil Giraffiid (G@eners
Appearance Pi rakons of the horn- -cores 408 HLL SDEGER 518
The Faunal relationships at ‘aie African
THE FossILizED GIRAFFIDS : : sie AA. Sivatherine sites. = ; » 526
Introduction : ! : ‘ =o ATA REFERENCES : : : ‘ 2a 40
GENERAL INTRODUCTION
With the discovery of several large fossilized giraffid teeth (Singer, 1954) and,
later, of a pair of horn cores (so-called ‘palmated antlers’) on the large exposed sand-
duned fossiliferous site on the farm ‘Elandsfontein’ near Hopefield (60 miles north-
west of Cape Town), it was found necessary to compare these specimens with others
found in Africa. During this study, it became obvious that a complete survey and
review of the numerous genera of fossil giraffids found on the continent was essential.
On surveying the literature, very little information on the extant Gzraffa camelopard-
alis was found. Consequently a large number of skeletons of the extant animal were
studied in the American Museum of Natural History (New York), in the Chicago
Natural History Museum, in the U.S. National Museum (Washington, D.C.), in
the Department of Comparative Zoology at Harvard University, in the Musée Royal
du Congo Belge (Tervuren, Brussels), and in the South African Museum, Cape
Town, so as to obtain statistically significant ranges of variation of intra-specific
1 Summary read at the 5th INQUA Congress, Barcelona in September, 1957.
2 Honorary Curator of Human Palaeontology, South African Museum, Cape Town.
3 Visiting Professor in Anatomy, University of Illinois, Chicago (1959-60).
375 SMITHSONIAN
NSTITUTION AUG 3 0 1960
VOL. XLV PART IV
376 ANNALS OF THE SOUTH AFRICAN MUSEUM
characters which would be of value in assessing the fossil specimens. This paper is
consequently divided into three* portions, the first dealing with observations on
the modern giraffes, the second containing observations on the fossil material
assembled in Cape Town or studied in the British Museum (Natural History), the
Muséum d’Histoire Naturelle (Paris), and the Coryndon Museum, Nairobi, while
the third section contains the discussion and conclusions.
SECTION I
SURVEY OF THE LIVING GIRAFFE
INTRODUCTION
In the literature numerous references are made to various subspecies of
G. camelopardalis. ‘These distinctions are based on skin colour pattern and on horn-
core differences (Lydekker, 1904). Owing to the fact that the skins of the extant
specimens studied were not preserved in most cases, no correlation with the skeletal
material was possible, but the study of the horn-cores (vide infra) indicated that no
substantial subspecific differences could be detected, and consequently it was con-
sidered reasonable to treat the material as a whole in a specific determination. As
a result of the observations made and the wide range of variation of the data obtained
for the whole group, it is suggested that the diagnostic criteria for subspecies of
G. camelopardalis should not be based on skin colour patterns alone as these may be
ecological variables. In this respect, it is interesting to observe in the list of extant
specimens examined (Chapter 2) that the deficiencies in the information regarding
the subspecies in Museum records are due to the fact that no accurate diagnosis was
possible on the basis of the skeleton alone when no skin was received.
CHAPTER I
GENERAL DESCRIPTION OF THE GIRAFFID DENTITION
(Applicable to modern giraffes and extinct giraffids.)
While almost all the Eocene Ungulata possess the full mammalian dentition
(Iz Ct P= M3 = 44), maxillary incisors and canines as well as anterior upper and
lower premolars and posterior molars have been lost in the course of the evolution of
the Giraffidae, so that all recorded fossil material already possess the formula of the
extant giraffes, viz. I$ C2 P2 M3 = 32. The mandibular incisors must have occluded
—as in the modern Pecora—against a hard elastic pad of the gum. Parallel to this
reduction in the number of teeth, a great development of the cheek-teeth is observed,
as well as a marked tendency to hypsodonty (less noticeable in the modern species),
both characters making the dental apparatus suitable for a herbivorous diet, which
* A fourth portion, the Appendix, was added later.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 377
requires a great amount of mastication. In the general pattern of the teeth, there is
no essential difference between fossil and extant Giraffidae, although quite a few
of the fossil genera have got much larger teeth as well as minor variations in structure.
As far as it can be judged from the extinct material available, the whole family is
characterized by a brachy- to hypsodont, selenolophodent type of molar. In all
specimens, the third lower molar has an accessory posterior lobe or talonid. The
somewhat cuboid crown of the molars presents on its occlusal surface—even before
any marked wear—a longitudinal fold, extending across the summit mesio-distally,
and a transverse cleft extending bucco-lingually, dividing the tooth into an anterior
and a posterior lobe or pillar. The external surface presents two vertical grooves
running from the grinding surface to the root, the buccal one being by far the
deepest in the lower molars, the lingual one in the upper; the less marked groove on
either surface is actually nothing more than the lateral depression resulting from the
formation of a mesostyle by an elevation of the cingulum (vide infra). Each of the
pillars of the molars are slightly angulated through their transverse axes to the
main longitudinal axis of the jaw so that the former axes are rotated anteriorly and
medially: the anterior portion or pillar of a particular molar lies more laterally
(buccally) than the posterior pillar of the preceding molar.
It is not essential here to present details and discussion about the origin of the
Ungulate tooth and its components, but a brief description of the structure of the
giraffid tooth is justified for the sake of clarity.
In earlier mammalian forms, trigone and talon distinguish between cutting and
crushing portions of the crown, but the trigone is lowered in Ungulates to the level
of the talon. Four main primary cusps may be recognized in the molars,’ while a
fifth one, derived from the cingulum, has probably been established very early as a
further crown cusp. These five cones (or conids in the lower molars), which are
elevations of the grinding surface of the tooth, eventually become separated from
one another by small secondary and intermediate cusps, conules or conulids. The
periphery of the base of the crown, or cingulum, encircles the neck of the tooth: it is
well or little developed, depending on a rather wide individual variation. Never-
theless it is nearly always present and commonly gives origin to peripheral cusps,
called styles or stylids. In this paper, cusps are named according to the tritubercular
theory of Osborn (1892), but this does not necessarily indicate that there is an
acceptance of all its broad phylogenetic implications. As this paper is exclusively
devoted to Giraffids, the main purpose of utilizing Osborn’s nomenclature is to
provide a clear description of morphological features. However, in this respect, the
suggestions of Arambourg (1947) are very helpful, because his amended nomen-
clature may be applied both to molars and premolars. Main cusps will therefore
be called cones or conids. ‘The medial (lingual) ones are, in the upper teeth, from front
to back (mesio-distal), the protocone and the hypocone, in the lower teeth the paraconid,
the metaconid and the entoconid (fig. 1). ‘The lateral (buccal) cusps are, respectively,
the paracone and the metacone, and the protoconid and the hypoconid. Secondary cusps
in the upper teeth are the intermediate paraconule and metaconule, derived from
elevations of the crown, and, in the lower, the hypoconulid. Other secondary cusps
378 ANNALS OF THE SOUTH AFRICAN MUSEUM
are also derived from elevations of the cingulum, viz. the lateral parastyle, mesostyle
and. metastyle, and the protostylid, ectostylid and hypostylid of upper and lower molars
respectively, while on the medial side are the protostyle, entostyle and hypostyle, and the
parastylid, metastylid and entostylid on the upper and lower teeth respectively.
In the Ungulate type of tooth, especially among Pecora, several cusps or styles
lose their individuality and fuse into crests or ridges or lophs. These are built up of
pas’) Pa MS UOT mts
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Fic. 1. Schematic representation of giraffid molar and premolar nomenclature
(after Arambourg, 1947).
A, upper molar. B, upper premolar. C, lower molar. D, lower premolar.
Upper jaw Lower jaw
Main cusps Internal 1. Erotocone ) 2 Paraconid Pad
2. Hypocone H Metaconid Md
2. Entoconid Ed
External 1. Paracone Pa Protoconid Pd
2. Metacone M Hypoconid Hd
Secondary cusps Median anterior Paraconule Pl
posterior Metaconule ml Hypoconulid hld
External I. parastyle pas protostylid psd
2. mesostyle ms ectostylid ecd
3. metastyle mts hypostylid hsd
Internal I. protostyle ps parastylid pasd
2. entostyle es metastylid msd.
3. hypostyle hs entostylid end
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 379
two or more cusps and are ordinarily very marked, so that protoconule and protocone,
metaconule and hypocone, parastyle, paracone, and mesostyle, and mesostyle, metacone and
metastyle (or corresponding conids and stylids in the lower tooth) are fused forming four
of these crescentic ridges or ‘crescents’, the former two being more curved, and the
concavity of the crescents facing the buccal or lingual surface of the tooth in the
upper and lower molar, respectively. This double-pillared and double-crescentic
crown is the typical simple ruminant pattern and determines the seleno-lophodont
type of Giraffid tooth.
Prolonged into the depth of the crown by their enamel coat, the two parallel
rows of cusps and their linking crests [the buccal and the lingual, i.e. the proto-
hypoloph (metaloph, Osborn, 1907) and ectoloph] build the side-walls of the infundibulum,
central pit or valley, poorly lined by cement (crusta petrosa) in Giraffids, which
becomes stained by the food and constitutes the so-called mark (Marken, Bohlin,
1926; marque, Arambourg, 1947). The znfundibulum is itself festooned along the mesio-
distal axis of each pillar, and is contiguous in the transverse cleft with the central pit
of the adjacent pillar before any advanced wear takes place. When unworn, the
crescentic ridges are very well marked. With the gradual wear of the pointed
enamel crests, however, the apex of each loph tends to be levelled mesio-distally
while the lophs broaden in the bucco-lingual axis. An increasing crescentic tract of
dentine shows between the two newly formed lips of enamel which progressively
separate from one another; at that time the znfundibulum or central pit narrows by
approximation of the buccal and lingual lips of the adjacent walls of the crescents.
In very worn teeth, the pit may be restricted to a simple ridge of enamel, circular
(island of enamel) or linear, which represents the actual fusion line of the side walls
of the infundibulum in the depth of the crown, or the enamel of the pit may even
disappear completely.
It has often been observed and mentioned (already by Owen, 1840-5) that in
the giraffe, the median convexity (costa) of the buccal surface of the anterior pillar
in upper teeth (inner or lingual surface, in lower teeth) is more prominent nearer the
occlusal end of the crown than nearer the base, while the vertical ridge of the meso-
style (-stylid) projects outwards from the surface of the tooth more than the median
costa of the metacone (or entoconid). ‘This feature is quite consistent throughout the
family, and is distinctly recognized in the fossil material from the Siwaliks, India, as
well as in the specimens presently studied (vide infra).
The upper molars differ from the lower molars in their general shape, the
breadth or transverse (bucco-lingual) diameter approximating or even surpassing
the length or antero-posterior (mesio-distal) measurement.
As in most artiodactyls, the premolars are unilobed. However, in the fossil
specimens the last lower premolar has a clear demarcation of the posterior portion
of the tooth.
The incisors describe a semicircle, although in most ruminants they are arranged
in a straight transverse line at the extremity of the jaw. The canines have been
profoundly modified in shape and position and resemble very much a fourth pair of
incisors; however their crown is larger, much more triangular and somewhat bilobed.
380 ANNALS OF THE SOUTH AFRICAN MUSEUM
All the teeth are covered by a coat of relatively very rugose enamel, both on the
extant and fossil specimens. This feature is characteristic of the group. The maxi-
mum rugosity is always observed on the medial surface of the protocone and hypocone, ©
and on the lateral surface of the protoconid and hypoconid.
Incisors and canines are implanted by a single root which slopes backwards
horizontally. The lower teeth have two roots, an anterior and a posterior one; all
the upper teeth have three roots, two lateral and one medial. However, both
anterior and posterior roots of the lower molars as well as the medial root of the
upper molars show a tendency of fusion of two fangs. They are broadened, flattened
from side-to-side or bucco-lingually, and present a median groove, the groove being
on the lingual surface in the upper, while in the lower the groove on one root faces
that on the other, the deeper groove being on the anterior root. ©
CHAPTER 2
LIST OF RECENT GIJRAFFA CAMELOPARDALIS STUDIED
The observations recorded below on the extant material were made on the
following specimens, for which the number, age, sex, origin and subspecies have
been extracted directly from the registers of the respective museums. Gaps in the
information below indicate deficiencies in the registers.
I. U.S. NATIONAL MUSEUM, WASHINGTON, D.C.
No. Subspecies Age Sex Origin
14411 Gc. camelopardalis 1@) — —
121010 G.c. rothschildi AG) M Lake Baringo, B.E.A.@
154033 G.c. capensis I M Matabeleland, Rhodesia.
155438 G.c. rothschildi A M Guas Ngishu Plains, B.E.A.
162016 G.c. tippelskirchi A — Kilima Kui Kapiti Plains, B.E.A.
162017 G.c. tippelskirchi A M Kilima Kui Kapiti Plains, B.E.A.
162018 G.c. tippelskircht A M Ulu Station, B.E.A.
162988 G.c. tippelskirchi A F Sotik, Gnaso Nyiro, B.E.A.
162989 G.c. tippelskirchi I F Sotik, Gnaso Nyiro, B.E.A.
163112 G.c. rothschildi®) I M Guas Ngishu Plateau, B.E.A.
G.c. tippelskirchi I M Sotik, Gnaso Nyiro, B.E.A.
163113 G.c. reticulata A — Gnaso Nyiro, B.E.A.
163312 G.c. rothschildi A — Guas Ngishu Plateau, B.E.A.
163324 G.c. reticulata A F N. Gnaso Nyiro, B.E.A.
182124 G.c. reticulata A —- Koga Water, B.E.A.
182125 G.c. reticulata A — Marsabit Road, B.E.A.
182192 G.c. reticulata A —- Lakiundu River, B.E.A.
200151 G.c. rothschildi A M Wasin Gisher Plateau, B.E.A.
251797 G.c. tippelskirchi A F Savanda, Dodoma, Tanganyika.
251798 G.c. tippelskirchi A — Dodoma, Tanganyika.
251799 G.c. tippelskirchi A — Mkata Plains, Tanganyika.
251800 — A — —
252549 a I M Sudan (Nat. Zool. Park).
252585 — I F Sudan (Nat. Zool. Park).
270594 G.c. reticulata A F — (Nat. Zool. Park).
279405 — g years M — (Nat. Zool. Park).
296145 — A — S.W.A.@), Gaucha (about 19° 47’ S.
20° 35’ E.).
299998 G.c. camelopardalis I M — (Nat. Zool. Park).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 381
II. CHICAGO NATURAL HISTORY MUSEUM
No. Subspecies Age Sex Origin
27475 oj I M Northern Uganda.
29515 — I ¥ Kenya.
32901 G.c. reticulata A F Abyssinia, Sidamo, Boram Border.
32902 — A F Abyssinia, Sidamo, Boram Border.
32904 G.c. reticulata I M Abyssinia, Sidamo, Boram Border.
32905 — I M Abyssinia, Sidamo, Boram Border.
34422 == A M Bechuanaland, Mababe Flats.
34423 _— A F Bechuanaland, Mababe Flats.
34424 — A F Bechuanaland, Mababe Flats.
34425 — A M Bechuanaland, Kalahari Desert,
Kwaai, Mokaba River.
34426 — A F Bechuanaland, Mababe Flats.
34427 — I F Bechuanaland, Mababe Flats.
34428 —_— I M Bechuanaland, Mababe Flats.
34429 — A F Bechuanaland, Mababe Flats.
34930 — A M Kenya.
53705 G.c. reticulata A M —
54251 — I M Tanganyika (Zoo).
xX, —_ A om ct
III. COMPARATIVE ZOOLOGY MUSEUM, HARVARD UNIVERSITY
8370 G.c. tippelskirchi A — East Africa.
837 (5) pee A ae —
83 72(5) — A aes a
14564 G.c. tippelskirchi A — B.E.A.
15698 — A — Sudan, Dinder River, near Kuka-
Dindu.
27137 — A — East Africa.
IV. AMERICAN MUSEUM OF NATURAL HISTORY, NEW YORK
20884 — I F _
24290 — A — Bechuanaland.
24291 — A = Bechuanaland.
24292 — A —- Bechuanaland.
24203 — A — Bechuanaland.
27675 — A — Kenya.
27752 G.c. rothschildi A — B.E.A.
27753 G.c. rothschildi I — B.E.A.
35536 se A M =
35628 — I — —
53543 G.c. congoensis A F Belgian Congo.
53544 G.c. congoensis I — Belgian Congo.
(Foetus)
53546 G.c. congoensis A F Belgian Congo.
53548 G.c. congoensis I M Belgian Congo.
53549 G.c. congoensis A F Belgian Congo.
54122 G.c. tippelskirchi I M B.E.A.
54123 G.c. tippelskirchi A M Northern Gnaso, Nyiro.
57675 = A = =
69403 — I — —
80146! (6) _ I M (New York Zoo)
7 months
80146? ©) Be I a ys
81820 — A — —
81821 — A — =
81822 — A — —_
81823 — A — —
81824 — A ~— —
81825 — I = —
81826 — I — i
382 ANNALS OF THE SOUTH AFRICAN MUSEUM
No. Subspecies Age Sex Origin
82001 — A M Kenya.
82002 — A F Kenya.
82003 — I — Kenya.
83458 — I F Bechuanaland, Mababe Flats.
83459 — I F Bechuanaland, Mababe Flats.
83460 — A M Bechuanaland, Mababe Flats.
83605 — I — Southern Rhodesia, east of Ngamo
Station.
99493 — A M Nubia (23 years in a zoo).
139695 7 A a a
139696 — A — —
144915 — A — (Central Park Zoo, N.Y.).
165051 G.c. capensis I M S.W.A., Etosha Pan, Farm Lom-
bard.
165052 G.c. capensis I F S.W.A., Etosha Pan, Farm Lom-
bard:
x, da A a sl
x, @ oe A w: aS
Xy (7) Fy A pe a,
V. SOUTH AFRICAN MUSEUM, CAPE TOWN
M. 245 = I — —
17176 — A — —
eds pa a a ae
VI. MUSEE ROYAL DU CONGO BELGE (TERVUREN, BRUSSELS)
R.G. 4947 G.c. congoensis A — Uele.
R.G. 4948 G.c. congoensis A — Uele.
R.G. 4949 © G.c. congoensis A M N.E. Uele; sources of Garamba
River.
R.G. 6342 G. A — Ufipa district.
R.G. 2128 G.c. schilbergsi A — B.E.A., Serengeti Plains.
(1) ‘I’ signifies immature; ‘A’ = adult.
(2) B.E.A. signifies British East Africa; S.W.A. = South West Africa.
(8) The same number has been given to the skull and the jaws. Although they obviously belong
to the same individual, the skull is labelled ‘rothschildi’ and the jaws ‘tippelskirchi’, and they have been
given different places of origin.
(4) This specimen has no Collection number. This indication is ours.
(5) Although skull and jaws have the same number they do not belong to the same individual.
(6) These two specimens have the same number. We refer to them as (1) and (2).
(7) These three specimens are not registered. We refer to them as X,, X, and X,.
CHAPTER 3
OBSERVATIONS ON TOOTH ERUPTION AND CRANIAL SUTURES
Among the specimens examined, a group of 32 immature individuals forms a
rather complete series of the various juvenile and adolescent stages of growth. In
describing this series, the sequence of tooth eruption and cranial suture synostosis,
and some data concerning the milk dentition and the growth pattern are particularly
emphasized.
From the information available, it is possible to extract the subspecies of Giraffa
camelopardalis in 12 cases: camelopardalis (2), rothschildi (2)*, tippelskirchi (2), congoensis
* The same number (U.S. Nat. Mus. 163112) has been given to the skull and the jaws. See
note (3) in legend in previous chapter.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 383
(2), reticulata (1) and capensis (3). As far as the sex is concerned, 10 specimens were
registered as female, 13 others as male. As regards their origin, it is known that
8 animals derive from British East Africa, 2 from Sudan, 2 from Ethiopia (Abyssinia),
2 from the Belgian Congo, 2 from Southern Rhodesia, 4 from Bechuanaland and 2
from South West Africa. Some specimens of unknown origin had been received from
zoological gardens or veterinary colleges. The ages extend from the foetal period
to the sub-adult stage. Information concerning the precise age of two specimens
were found in the Museum registers, viz., specimens 299998 and 801461 which are
5 months 26 days and 7 months old respectively; unfortunately the latter animals
were born, reared and then died in zoos. It is quite likely that conditions of captivity
affected the growth tempo, and data concerning these must be evaluated with care
before using them as age standards.
A. TooTH ERUPTION
In the group of immature animals 826 teeth were available for study (table 1).
In this number are also included the sockets of missing teeth. Furthermore, although
Paar Oenmrion I, I, I, C PP? P, P? P, P* P, M! M, M? M, M? M,
(411) P2eior io oo TG. If tO! 1G To fC ~ho. 5a = 44 Ast M96" 26
Mitk DENTITION is Io Ig c DM? DM, DM? DM, DM* DM,
(415) 32 36 440C 4842 42 42 420 45048
TABLE 1. Number of teeth examined in the immature giraffes.
some of the teeth were either broken or incompletely erupted, valuable information
was gained from observations on either their roots or their sockets. Accurate measure-
ments could be made on 331 perfectly preserved milk teeth. The stage of eruption
of the dentitions is indicated in table 2 and figure 2.
EruptTinc TooTuH STAGE I STAGE 2 STAGE 3 STAGE 4 ABSOLUTE AGE
DM3 53544 Foetus
Mi 163112 299998 I44il 6 months
34428 82003
80146?
83459
M2 252549 162989 7 months
252585 53548
801461 54122
54251 81826
M3—I1 165052 29515 83605 81825
20884
P4, P3, Pa 35628 69403
27753
Ig 154033 34427
165051
C 27475 32905
83458
TABLE 2. Stage of eruption of individual teeth. The appearance of the tooth at the surface of the
bony alveolus is stage 1, while stage 4 indicates complete eruption of the crown. Stages 2 and 3 are
equally spaced intermediate phases.
384 ANNALS OF THE SOUTH AFRICAN MUSEUM
The observations on the state of eruption and wear lead to the following
conclusions:
1. For both the milk and permanent dentitions (with the exception of DM4,
vide infra), the upper teeth erupt slightly before the corresponding lower teeth. The
priority of eruption of a particular upper tooth was observed in 16 of the 18 cases
(from 13 different individuals) where the difference between the time of eruption
of a tooth in the upper and of a corresponding time in the lower jaw (here called
‘asynchronism’) could be traced, namely, for M1—80146?, 82003, 83459; for M2—
81826, 53548 and 54251; for M3—81825, 83605, 165052 and 29515; for P2—27753
and 35628; for P3—27753, 35628 and 83605; and for P4—30628. In only two cases
the lower tooth has erupted shortly before the corresponding upper tooth, viz. in
69403 (P,) and 299998 (M,). In no case, however, is the asynchronism too marked:
in the above 18 cases, the more precocious tooth is never fully developed before the
lower one starts erupting. Four degrees of actual eruption are defined from the first
appearance of the tooth on the alveolar surface (stage 1) to its final development
(stage 4); two intermediate stages (stages 2 and 3) of progressive eruption are inter-
posed. The state of development prior to any eruption is termed stage o. In 17 of
the 18 cases the degree of asynchronism does not exceed one unit (i.e. o—1, 1—2,
2—3, 3—4), while in only one case, viz. the P3 of 35628, is the difference between
the stage of eruption of the upper and the lower tooth 2 units (1—3). It is not easy
to evaluate the absolute duration (in terms of days, weeks, etc.) of the asynchronism.
Eruption may be a rather rapid process because in one particular case (35628), the
milk molar DM* is still fixed on the summit of a fully erupted P*. No conclusions
can be drawn whether there is lack of asynchronism of eruption between right and
left corresponding teeth in a maxilla or mandible. The evidence suggests that they
erupt synchronously (fig. 2).
2. The complete milk dentition is in situ and is already functional before the
first permanent tooth (M?) erupts, e.g. 80146, 82003, 83459, 163112, 299998, 14411
and 34428. The first two incisors i, and i, probably precede the appearance of the
first milk molar. Only one specimen of the collection (53544) is at such an early
stage, but because it is rather badly preserved it is not possible to draw a definite
conclusion. The first two incisors are perfectly formed and fully erupted before
DM, begins to come through, but the evidence is not sufficient to ascertain the
relationships of i, and the canine, although specimen 32904 suggests that the latter
appears immediately after all the milk molars have erupted.
3. The milk molars erupt in rather regular succession, posterior to anterior in
the mandible, DM, preceding DM3, which in turn precedes DM,, as in specimen
801467 and 53544. Furthermore, the latter is still a foetus and indicates that the
whole milk dentition is formed before parturition. The succession is reversed for the
maxilla, as judged from specimen 80146? where the eruption starts with DM?. On
the other hand the six upper and lower milk molars seem to erupt simultaneously
in specimen 32904, so that one cannot generalize dogmatically on the basis of this
small series.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 385
4. The permanent molars of both upper and lower jaws erupt in the same
order, i.e. from anterior to posterior: M1 precedes Mg@ in 14 cases (53548, 801461,
801467, 81826, 82003, 83459, 14411, 163112, 252549, 252585, 299998, 162989, 34428
SS S5SS
a Sa <
= — (av Oo ont "”
SS SS Ge aa Gm A! a
14411 o* 0° 0° 0% o® 0° o° o°
20884 er eee ew le ee wef | e® of 0" 6”
27475 e® 0° 0 0° oe |e® 0° 0° 0° «0° 0° 0 0° of -°
27753 oe eo 0° 0° of of le” of 0° 0&8 of 0°
29515 e® o® 0° 0° 0% e° o° 0° 0° 0% 0° 0° 0° o° Ooh 2
32904 O° fen orn) OF |e" 16°. 0° oo e°
32905 of of 0° 0° 0° | 0° 0° 0° 0° 0° |e 0° 0° 0° 0°
34427 °° o* ef of ef o°/ 0° o° «° «° *ice° « © ce 0°
34428 o* 0° 0° 0° 0° | 0° e° 0° oe o°| -
35628 e° 0° 0° eo 0° 0° of ef le” 0° co O° 0°
53544 e* e° °° °° O°
53548 o*|e* of 0° 0 0° 0° of o® 0°
54122 " ef) e® 0° 0 0 eo / 0° o° 0° o°
54251 @7 e° e° e° oe" 0° of @° @°\\e° of oc 6°
69403 ee eo” 0° 0° 0° 0° | 0° 0° 0° 0° oe \e* 0° 0° o°
go14e! Stor res i. |eee™ |. oe
801462 Beta er se (e* lates ie) 66) co =
81825 ef of 0° | 0° 0° 0° 0° of |e® 0° 0° 0° 0 | 0° 0°
81826 of 0° 0° 0° 0°) 0° 0° © 0° 0° | 0 0° 0° 0°
82003 of of of 0° of 0° 0° of 0° 0% [0° .”
83458 e° eo” 0° o° 0° 0° | 0° o° 0° 0° wo \0° o® eo of .*
83459 eet ce" ete lee ete” eo" 6? | 0° =
83605 e° o° of 0° o® | 0° 0° 0° 0° 0° | 0° 0° 0° o = {0° 0°
154033 e° e° ef 0° 0° of 0%) 0° o° 0° of o° |e* o° 0° of -°
162989 o® ef 0° of of |e® 0° of o* 0° (0° 0° o° o°
163112 e® ef oe of 0° |e® o® 0 0° o° |.
165051 e° °° eo” 0° 0° 0° 0° |6® 0° 0° 0° eo je% 0° of 0° »°
165052 e® o° 0° oe of \0e° eo 0° o° 0° | 0° o° o° of
252549 oF 2° 0° 0° 00° 0° 0° 0° of |0® e® 0° 0°
252585 eo” 0° 0° 0° 0° (0° 0° 0° 0o° of 0° 0° 0% 0°
299998 eo” 0° 0° 0° ev le” 0° 0 @° 0° |0° o°
Fic. 2. Stages of eruption of individual modern giraffe dentitions
(upper and lower, left and right sides).
.—Stager. © —Stage2. © — Stage3. @ — Stage 4.
and 54251), and M2 precedes M3 in 19 individuals (20884, 53548, 54122, 80146},
81825, 81826, 83605, 165052, 252549, 252585, 162989, 54251 and 29515). No
exception to this order was found.
5. The replacement teeth of the milk molars, the premolars, usually appear
386 ANNALS OF THE SOUTH AFRICAN MUSEUM
immediately after the eruption of the last permanent molar, i.e. M3, and never
before it. In one case, however, M3 (83605) is still in the process of eruption (stage 3)
when P39, the first actual premolar to erupt, already cuts through the alveolar border.
The sequence of appearance of the premolars is less rigid than the eruption order of
the molars. In the four cases where the observation was possible (25628, 69403,
27753 and 165051), P2 is later than P3; in a fifth case (83458) P2 is bilaterally
absent in the mandible, and this might be considered as an extreme of the normal
late eruption of P2. In quite a few adult specimens, the same congenital absence
of P2 was observed (e.g. 83460, lower left), or alternatively, there is a marked
reduction of P2 where the root is abnormal (single) and the body slightly shorter
(A—P) than usual (83460, 34423 and 34424). While P4 precedes P3 in two cases
(27753 and 35628), P3 erupts before P4 in one specimen (83605). Consequently,
it may be concluded that the premolar eruption sequence is variable, but that the
whole process of eruption must be rather brief. These small individual differences
have probably no or little functional influence or significance.
6. The permanent incisors erupt at the same time as the molars and premolars.
In this respect, useful information is provided by 13 mandibles. I, starts erupting
during the eruption and maturation of M, (83605), and it is always present when
M, has completed its growth (27753, 35628, 69403, 81825, 83458, 83605 and 165051).
It might erupt slightly before M3 (81825) ; usually, however, it does so immediately
after (83605 and 165052). I, appears later and never erupts before I, or before the
full development of all the molars and premolars (27753 and 35628); in both cases,
for example, the milk incisor i, is still in situ when the last erupting premolar (P,)
comes through. On the other hand, specimen 69403 has a perfectly developed I,
while P, is still erupting, which would suggest that both teeth erupt practically
simultaneously, unless one considers it as an abnormal delay in P, eruption (it being
the only recorded case where the upper tooth is slower than the corresponding lower
one, and P, is in fact perfectly and completely developed). I, can be found, though
still in eruption, on three specimens (83458, 154053 and 165051) which already have
their two other incisors and all their molars and premolars. I, is still lacking in four
other individuals which have all their premolars (27753, 35628, 69403 and 83605).
Consequently, it indicates that the eruption of the lateral incisor I, occurs after that
of all the other permanent teeth, except the canine.
7. The permanent canine was found in only three specimens of the immature
collection (83458, 32905 and 27475). In one of these, it erupts directly under the
milk canine which is still in situ, all the other permanent teeth being perfectly
developed. This animal died during the transition from a sub-adult to an adult
stage as is indicated by the fact that the canine is the last tooth to complete the
dentition of the giraffe.
Figure 3 illustrates a schematic and summarized view of the above information
as well as an attempt to determine the absolute age of tooth eruption in modern
giraffes (data completing and correcting Owen, 1840-5, and 1849, as well as
Lankaster, 1907).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 387
B. CRANIAL SUTURE AND TOOTH ERUPTION
In the same group of immature giraffes observations were made on eight sutures:
fronto-sagittal, fronto-parietal, inter-parietal, temporo-parietal, temporo-occipital,
DAYS | WEEKS MONTHS YEARS
foetus Or AS . 3 4 y 3. ie Sta 4% Sb 5) 6 6/7
ii ee EEE EE EEEE——————e——— ee
i2 a nly
M1 a ae ee ee Se ee ee ee
EE a Re Es ee PUREE «ae PDE ea ee a
M3 ——————————————— Ee eee
I1 a a
P4 en Ea ee Pn Se SONS aR PS Yo SURES SEED EO
P3 Se ER ee ee eS
rae TOLTY Wl Ry fe) Pie 9 OD naecpaleubete ele oe Se So a
12 Le a 2 ee ee
I3
sir reat ey Pt AS Ly me No) fea ee Ql lee =
Fic. 3. Schematization of the order of eruption of teeth in Giraffa camelopardalis
and a determination of the absolute age of tooth eruption.
@ — Teeth deeply embedded in bony alveolus (Stage 0).
1 — Begins to erupt through bony alveolar margin (Stage 1).
fa] — Just penetrating through bony margin (Stage 2).
§) — Crown projecting beyond bony margin (Stage 3).
& — Crown entirely erupted (Stage 4).
x — Tooth fallen out.
parieto-occipital, exoccipital-supra-occipital and palatal. ‘Three different degrees of
synostosis were determined: the bones may be in contact without any trace of
closure (0), the suture may be half (1) or completely (2) closed (table 3); see p. 388.
The main conclusions of these observations and of their correlation with the
tooth eruption order (fig. 4) are:
1. The inter-parietal suture closes first, rather early, probably at the time of
eruption of M1, and is immediately followed by the closing of the parieto-occipital
suture. The examination of these sutures was possible in only 20 specimens, but in
only one case (83459) the parieto-occipital suture had not started to close, although
M1 was already erupting (stage 3) and the closure of the inter-parietal suture was
slightly delayed (half-closed), but this may be an individual variation and does not
affect the general observations.
2. The occipital suture is the next to be fused. It is always closed when P3
erupts; in fact there seems to be a definite tendency to fuse even earlier, namely,
during the eruption of the molars, as seen in 34428, 801461, 20884, 29515 and 83605.
388 ANNALS OF THE SOUTH AFRICAN MUSEUM
Exoccipital-
Specimen Fronto- Fronto- Inter- Parieto- — Parieto- §Temporo- =‘ Supra- Palatal
Sagittal — Parietal Parietal Temporal Occipital Occipital Occipital
32904. ) oO fo) fo) O o (0) o
53544 O oO 0) ) O ) O co)
34.428 2 2 2 2 2 O 2 0)
82003 oO o 2 fe) I oO oO e)
83459 Oo o I oO o co) oO o
801461 ) fo) 2 fc) 2 o 2 O
54251 oO Oo 2 (a) 2 Oo Oo oO
54122 ) O 2 (a) 2 oO O fo)
81826 O ) 2 O I fo) fo) oO
165052 I ) 2 Co) 2 Oo O fc)
20884. O e) 2 I 2 o I oO
29515 ? O 2 O 2 o 2 (a)
83605 O oO 2 o 2 oO I oO
81825 oO (a) 2 (e) 2 oO oO oO
35628 I 2 2 I 2 ) 2 o
27753 I O 2 I 2 fC) 2 I
69403 oO oO 2 (0) 2 oO 2 (a)
165051 2 oO 2 oO 2 O 2 oO
34.427 2 ) 2 (0) 2 ) 2 oO
27475 2 ) 2 o 2 oO 2 o
83458 I I 2 Ce) 2 (0) 2 e)
32905 2 2 2 O 2 oO 2 )
TABLE 3. Cranial suture closure in immature giraffes. The skulls have been graded according to
their dental age (see text, and fig. 4 where the same information on the degree of suture closure is
correlated with the tooth eruption order).
oO
=}
oO = om U vD U0 x< & =< <= o (o) (ws) ° - = = e
G5 NB 21ND PANO UNESH lint ROY pea ye armen Shi te =
De) wo +s z=
——--0 ° ° ° ° Fronto -sag.
° ° ° Fronto-pariet
(nter parietal
Parjeto-temp.
Temp.-occipit.
Parieto-occip.
Occipital
° Palatal
Fic. 4. The relationships between cranial suture closure and tooth eruption order
in Giraffa camelopardalis.
© : Suture fused, at least at age of the tooth eruption stage.
—O© : From the stage of eruption of this tooth onwards, all specimens found with the suture
fused, i.e. in one specimen the occipital suture fused at age of eruption of Mr, in
another at age of eruption of Mg, etc....
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 389
3. The fronto-sagittal suture is always completely fused when the lateral incisor
erupts, although again there is a tendency to fuse earlier (at the time of eruption of
M3 or of the premolars).
4. The fronto-parietal suture fuses at the end of the sub-adult period of growth,
but, as a general rule, it is not completely fused before completion of growth.
5. The temporo-occipital and the palatal sutures close much later, during
adult life, although the latter has been observed partially fused in one case (27753)
at the eruption of P2. It has already been mentioned that late synostosis or retarda-
tion of closure may occur as an individual variation, and similarly, an acceleration
has been observed in specimen 34428, in which, at the time of eruption of M1
(stage 1), all the sutures normally fusing during juvenile or sub-adult life are already
completely closed.
C. SKULL GROWTH AND CRANIAL MEASUREMENTS
From a few cranial measurements (table 4), it is possible to draw some con-
clusions about growth patterns in modern giraffes. First of all, the ratio fronto-
sagittal length/basion-nasal length is practically constant, the range of variation
being relatively very small (31-6—36-9) and not altering with the growth gradient.
Similarly, the ratio basion-palatal length/palato-nasal length is practically constant,
indicating that both elements of the skull length (cerebral and facial) grow at a
rather parallel tempo.
Maxillary
Specimens from American Biorbital Basion- Palatal- breadth Frontal
Museum of Natural History diameter palatal nasal opposite M+ length
53544 IIT 78
82003 178 139 162 104 III
801461 175 112
83549 180 138 210 106 Lay
80146? 189 163 124 124
53548 217 183 260 127, 149
54122 203 129 139
20884 183 165 262 119 143
165052 243 183 207 140 156
81825 251 208 311 145 Heit
83605 235 194 304 139 171
35628 215 180 305 118 169
27753 255 200 249? 138 176
81826 209 169 133 154
165051 276 219 366 152 IQI
83458 268 209 142 174.
69403 276 207 336 152 177
Specimens from U.S.
National Museum
163112 186 133 208 107 120
14411 162 128 IQI 106 Rey
252549 E73 146 231 III 121
252585 184. 142 203 III 124
299998 177 147 203 113 123
162989 226 170 268 130 143
154033 268 570 149 QI
SL eEEEEEEEEEES Selene
TABLE 4. Cranial measurements (mm.) of immature modern giraffes. (Basion-nasion distance
obtained by addition of basion-palatal and palatal-nasal measurements.)
390 ANNALS OF THE SOUTH AFRICAN MUSEUM
Comparing further the growth of the fronto-sagittal length and of the biorbital
breadth respectively, one sees that parallel to the increasing length of the skull,
there is a very slight reduction of the biorbital breadth, and a more marked relative
decrease of the maxillary breadth (fig. 5). Another expression of the same trend is
that with the increasing absolute biorbital breadth, there is a decrease in the ratio
maxillary/biorbital breadth. Thus during the juvenile and adolescent growth
periods of giraffes, the upper portions of the skull, both facial and cerebral, develop
at a quicker tempo than the lower part of the face, the breadth of which decreases
relative to the biorbital breadth and the frontal length.
Max. x 100
Fr.-sag.
Fr.-sag.x100 Ms z
Biorbit. 5 ¢ - 100 %
oe e
. 95
75 Uo _
. 90
70 Eas
. 80
Bet
65 e e @
C) e -
° . 70
e@ : °
- 65
60 . °
Fronto-sag._,70 80 90 100 "0 120 130 140 150 160 170 180 190 210 210
(mm)
Fic. 5. Comparison of growth of the fronto-sagittal length and of the biorbital breadth.
CHAPTER 4
VARIATIONS OF THE CUSPS OF THE TOOTH CROWN
The variations of cusp-development have been studied in the dentition of the
extant giraffe. A large number of teeth have been surveyed (table 5). As regards
the cusp-pattern, there is very little difference between right and left corresponding
teeth in the same jaw: therefore the observations summarized here will concern only
one of each pair of teeth. In the following table, the numbers in brackets indicate
the actual number of individual pairs of teeth, while the non-bracketed numbers
indicate the actual number of teeth available.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 391
Po P3 PA. Mi Ma2 M3
UPPER 117 120 116 138 130 121
(59) (60) (58) (69) (65) (61)
LOwER 109 124 Br 143 135 124
(55) (62) (64) (72) (68) (61)
TABLE 5
First, the development of the main (crown-) cusps, cones or conids, and secondly
that of the secondary cusps, lateral styles or stylids, are dealt with. In each section,
the molars will be described first, and then the premolars.
Main Cusps
1. Molars. The four main cones and the five conids of all the molars surveyed
show no significant variability. In the upper molars, there is a rather equal develop-
ment of proto- and hypocone on the lingual side, while on the buccal aspect, the
paracone is regularly, and probably typically, more developed than the metacone of
the same tooth. This is applicable to all true molars, although the vertical costa of
the paracone seems to be more strongly built in M? than in M3, and still more in M! .
than in M?.
In the lower molars, the fused para-metaconid on the lingual surface of the
anterior pillar is slightly more developed than the corresponding entoconid on the
posterior pillar, but the difference does not show as markedly as in the opposing
upper tooth. The talonid is always very well defined in M,: its A—P occlusal length,
measured in 78 cases, is 23°3% (=: -31) of the total mesio-distal length of the tooth
(range of variation: 16:5-30°8; o = 2:70).
2. Premolars. The premolars show the typical main cone pattern: the unilobed
upper premolars have three cusps—the frotocone is on the lingual side, and the para-
cone and metacone on the buccal side are fused into one single enamel crest which
terminates mesio-buccally in the parastyle. In the lower premolars the more compli-
cated but typical features of the giraffid tooth are always recognizable: paraconid and
metaconid are fused into one single mesio-lingual ridge on P,, but they are more
isolated on the anterior premolars when the paraconid is much smaller, while in P,
it is restricted to an ill-defined formation on the outer (buccal) border of the crown.
The entoconid is always very well defined and linked with the protoconid on the buccal
aspect by means of a strong ridge of enamel running outwards (buccally) and
forwards (mesially) across the horizontal surface of the tooth The hypoconid on the
buccal aspect is always markedly separated from the protoconid by a well-defined
vertical furrow which, however, decreases from the grinding surface downwards
and disappears before reaching the bulging of the crown well above the margin of
the crown-root junction
SECONDARY CUSPS
The observations indicate that there is little variation in the main cusp pattern
of the modern giraffe teeth. In contrast, the secondary cusps show a marked degree
392 ANNALS OF THE SOUTH AFRICAN MUSEUM
of individual variation, some of the most characteristic expressions of which will be
described below. No emphasis will be placed on the median anterior or posterior
cusps, para- and metaconule respectively, because it is almost impossible to distinguish
these formations in many cases, as the demarcation between them and the proto-
or hypocone is hardly noticeable along the proto or the hypo- (meta-) loph.
Vertical elevation of the cingulum to form styles or stylids on the peripheral
enamel coat of the tooth is common. Various features can be observed which, for
the sake of clarity, are classified here into two main types: (a) the ridge-type and (b)
the column-type. Variations of both types are quite numerous.
(a) the thin ridge of enamel may appear either as a straight or a notched crest, and
may even have a comb-like appearance distally detached from the crown
(fig. 6).
(a) Ridge pe
i ii il
(b) Column
ra) / \
i li ii
Fic. 6. Types of styles or stylids developing from the
cingulum in Giraffa camelopardalis.
(b) The column type presents as a better defined formation rising from the cingulum
or the enamel coat either as a small tubercle or a developed narrow column or
pillar. In some cases this column appears to be truncated and flanked on both
sides by smaller tuberosities.
The column type secondary cusps are always found on molars and premolars,
anterior and posterior to the buccal paracone and metacone, and to the lingual para-
metaconid and entoconid of upper and lower teeth respectively, i.e. parastyle and
-stylid*, mesostyle and metastylid, metastyle and entostylid. The parastyle and metastyle in
the upper premolars, the parastyle and mesostyle in the upper molars, the entostylid in
the lower premolars, and the metastylid and the entostylid in the lower molars are
typically the best defined and the least variable of these lateral (buccal upper or
lingual lower) secondary cusps.
On the other hand, the lingual surface of the upper teeth and the buccal and
posterior surfaces of the lower teeth show a very marked variation in their styles
* The column-like parastylid however is usually rather poorly developed. In a few cases, it was
even observed being replaced by a more ill-defined crest-like stylid. This latter feature is shown in
one P,, two Py, two M,, five M, and five M3.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 393
(stylids). Indeed, in this region, both types of cusps occur, but it should be noted
that while the ridge-type has been observed on both premolars and molars, the
column-type has only been observed on molars. Furthermore, while ridges may
represent the protostyle (-stylid), the entostyle (-stylid), the hypostyle, the ectostylid or the
hypoconulid, the column type secondary cusps recorded are ALL (on these surfaces)
entostyles (-stylids) and ectostylids. Such formations on the lingual surface of the upper
teeth and on the buccal and posterior surfaces of the lower teeth have been observed
in 294 cases of the 754 pairs of teeth studied; in six cases, however, two styles were
observed on the same tooth, so that the actual proportion of teeth affected by these
secondary styles (stylids) is 38:2°%. The following table illustrates the distribution of
these styles among the types of teeth; the figures in italics represent the column-type
styles, while the non-italicized ones represent the ridge-type cusps.
P2 P3 P4 M1 Ma2 M3
PROTOSTYLE I I I 5 5 4
(1°7%) (1°7%) (1°7%) (7°2%) (7°7%) (6°5%)
7 7 6 I
ENTOSTYLE (11°9%) (11-°7%) (10°3%) (1°4%)
Cag 25 33
(39°2%) (38-4%) (54°1%)
HyYPosTyLE 4 5 5 I
(6-8%) (83%) (8-6%) (16%)
PROTOSTYLID 2 4 2 20 18 17
(3°6%) (6:5%) (3°1%) (27°8%) (26°5%) (27°9%)
EcTOsTYLID I I 2 2
(16%) (1°4%) (2:9%) (3°3%)
43 20 18
(59°7%) (29°4%) (29°5%)
HyYPOCONULID I I I I I
(16%) (16%) (1°4%) (1°4%) (16%)
ENTOSTYLID 2
(3°3%)
TasLe 6. Distribution of secondary cusps in extant giraffe teeth. (Bracketed figures indicate per-
centage of pairs of teeth available.)
The above figures show clearly the degree of variation of secondary cusp
formation in the extant giraffe dentition. It indicates that the tendency for such
additional features is much more developed in the molars than in the premolars, and
it is probably more in the lower than in the upper teeth. In the upper molars the
tendency increases in a posterior direction, while in the lower teeth M, seems to be
the most affected, with an average of o-g of a cusp per tooth, the corresponding
average for M, and for M, being 0-6. Furthermore, 7 ectostylids were routinely
recorded because of their big size: all of them were observed on M,. Entostyles and
corresponding ectostylids are the most frequently occurring secondary cusps: they
represent 36-1% and 29:6% of the total respectively.
394 ANNALS OF THE SOUTH AFRICAN MUSEUM
CHAPTER 5
DIMENSIONS OF THE TEETH
A. ApuLT DENTITION
From the maximum antero-posterior (mesio-distal) and transverse dimensions
of each tooth, the ranges of variation have been established, as well as the mean
and standard error (M + s.e.), the standard deviation (c), and the coefficient of
variation (V) (table 7). The data, as well as the indices, have also been expressed
EXTANT ADULTS
A -P = Tooth-length |
5 15 25 35 45mm
Fic. 7. The ranges of variation of the A—P length of the permanent teeth in Giraffa camelopardalis.
{| = the mean value.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 395
Giraffa camelopardalis
EXTANT ADULTS
Transv.
4 8 12 16 205 924 128 32 36> 407mm
Fic. 8. The ranges of variation of the transverse diameter
(breadth) of the permanent dentition.
| = mean value.
in graphic form (figs. 7, 8 and g). It may be observed from the mean and from the
relative dimensions of the teeth (fig. 10), that there is a decreasing gradation of size
for the postcanine teeth from back to front, except for M2 which is in general longer
(A-P) in the upper jaw, and broader in both jaws, than M3. The decrease in size
of the teeth in both upper and lower jaw is gradual, except for P2, which is relatively
396 ANNALS OF THE SOUTH AFRICAN MUSEUM
much smaller than P3 (more in the lower than in the upper). In the family Giraf-
fidae, P1 has disappeared, while the unusual decrease in relative size observed in
Pa, linked with the fact that in some specimens it is also absent (see chapter 3), may
suggest an eventual disappearance of Pa.
elralte) ec pmelonaiaelie
ULTS
Nt ae Tr: A-P Index
30. 40 50° GO. 7085580 190), 100", NO 120 G0. 40 Zo
Fic. 9. The ranges of variation of the transverse/A—P index of
the permanent dentition.
{ = mean value.
Both the premolars and the molars of the upper jaw tend towards a square
shape, the breadth being normally even greater than the A—P length (fig. 10). In
table 7 (and fig. 9) this is indicated by the fact that the mean of the transverse/A—P
397
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
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UuOT]eLIe A, ae UOT}eLIV A ean uOTIeLIE A sas
A > jo osury as aN | N A 4 jo osury eae N A jo osury mae | N p00],
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398 ANNALS OF THE SOUTH AFRICAN MUSEUM
index is greater than 100. On the other hand, in the lower jaw, the A-—P length is
always greater than the breadth, so that the teeth tend to be rectangular in their
longitudinal axis.
ae Giraffa Giraffa_camelopardalis
304 x 32.6 31.7x 33.8 267x309 222x296 226x281 207x255
5 263x242
iJ 3x24. '
a. sxe te 3x77
ra
ane
@&
= 2
~ = DM4 DM3 DM2 Cc ig in ty
JS €
& S
= ae eae
18.5 x66
206x137 x93
329 x 171
oc
= 381 x 23.3 303x236 283x222 245x217 220x205 178x155
= ste a ae ee
= 253 x 102
M3 M2 M1 PS) PSY Spo c |. Bee
Fic. 10. Graphic representation of the relative ranges of dimensions (A—P length and breadth)
of adult and milk (immature) dentitions. Mean dimensions are indicated.
Although similar in breadth to the incisors, the canine is approximately twice
the length of each incisor, probably because of its bilobate nature. The incisors
show a decreasing length (A—P) from front to back, although they have approximately
the same breadth.
Index N Range of Variation M s Vv
p2
Length = 122 73°8— 109°2 91°940°527 5°86 6:37
P2
Breadth pe 106 74°0—104°8 91°28-+0°429 4°38 4°79
tnd Transv./A—P P? 3 8-1 3 ee Bee
nde ScanEnEienenraraietiemtien 10 ‘O— 131° : 0°841 : :
i Transv./A—P P% ? : ate cae : ? ?
P3
Length =i 121 84°9—124°0 101°60+0°572 6°30 6-20
ps |
Breadth par 109 79°6—105°9 95°02-+0'373 3°88 4:08
Ind Transv./A—P P? 3 6 PAN aye 6-8 a
ndex = 4—————___—__ 4 oye . ‘19-0: ‘B80 :
Transv./A—P P4 Me ge ay os 4 ae q
P
Length ar 109 56-2— 94°8 80:40+0°746 7°76 9°65
3
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 399
Index N Range of Variation M S V
P,
Breadth = 110 57°7— 98:8 76-70-+0°665 6-92 9°02
3
“2 Transv./A-P P, : :
ndex Tea eas 109 74°2—130°0 95°84+1:042 10°84 11°31
P
Length = 123 77°8— 103°1 90°23-+0°396 4°36 4°83
a
P,
Breadth = 116 68-8—105:0 93°17-0°570 6-10 6°54
4
meee Es 8-8—128 64-£0-78 8-6 8
ndex aeeiney. ALP SP) I2!I 7 FS MELO! 103 40°7 I O 29
Be la
Length Ve 127 80°5— 101°6 93°12+0°348 3°90 4°18
Ml!
Breadth ME iO 81:3— 98:8 91°44-+0°320 3°52 3°84
= Transv./A—P M1}! ee P 3 3
ndex The eae 120 4°3—108°9 QS°12+0°412 4°50 4°5
M2
Length a 122 84°6—140°8 104°13+0-°665 7°32 7°02
M2 ;
Breadth Me 116 92°2—119°0 104°12+0°413 4°41 4°23
ret Transv./A—P M2? 5 oe Ewe 6
e ——— EEE 7 . . ° A ° °
ndex Transv./A-P M3 114 C71) Wao) 100°15+ 0°61 5 55
M,
Length Ni. 135 74°8— 106-0 93°7640°415 4°82 5°14
2
M,
Breadth “Ta 134 83:9—101°8 94°38-+0°276 3°18 3°36
2
Ind Transv./A—P M, ae 3 ie 3
nae SS SE Rr eae I “Q—I ° ° ° ° ° 2
x Transv./A-P M, 33 9—127 100°99= 0°520 on 59
M
Length aa 125 69°8— 95°4 _— 79°58-+0°421 4°68 5°88
3
M,
Breadth a 120 87-1 —133°4 101°34+0°372 4.06 400
3
os Transv./A—P M, ReCaEAG 6 6-68
eee 109°0— 150° 127° oO: : .
et Transv./A—P M, ue 2 oma? ales ane 2°
. Ps
Breadth il 108 73°6—106-2 91 -03-++0°469 4°84 5°31
Pt
Breadth Mi 108 83°0—109°5 95°740°458 4°72 4°93
P,
Length —— 124 69:0— 89-6 80-96-0388 4°31 5°32
M,
B
Breadth a 121 76-6— 104-2 92°62+0°485 5°34 5°76
2
TABLE 8. Dental Index for lengths and breadths, as well as the index for the transv./A—P ratios of
successive permanent teeth of G. camelopardalis.
400 ANNALS OF THE SOUTH AFRICAN MUSEUM
The ratios of the length and of the breadth of successive postcanine teeth
(P2/P3, P3/P4, M1/M2, M2/Ms3) are expressed as the ‘Dental Index’ for premolars
and molars. For both their lengths and breadths, the range of variation, mean,
standard error, standard deviation and coefficient of variation were determined
(table 8, fig. 11). In addition, the ratios of the transverse/A—P dimensions have been
compared and expressed as an Index. This data has proved particularly valuable
in assisting the authors to determine the diagnosis of isolated fossil teeth. In this
respect, the ratios of the breadths of P?/M!, P4/M}, P IM and of lengths of P,/M,
have also been helpful (table 8).
B. DEctipuous DENTITION
The measurements of 331 milk-teeth are summarized in table 9 and figs. 10,
12. It is clear that the front teeth (incisors and canines) and the first molar, both in
the mandible and the maxilla, have the largest coefficient of variation, while the
postcanine teeth show much less variation of both length and transverse breadth.
In most of the cases, the breadth seems to be slightly less variable than the length.
A-P (mm.) Transverse (mm.)
Tooth
Range of Range of
N ~— Variation M o Vv N_ Variation M a Vv
1, 23 «9th —13°5) «-11°53+°225 1°08 9:36] 22 6:0— 8-2 +7-48+-127 o-60 8-02
le 24 7°2—13°5 «11°27+°308 «61°51 13°399| 23 5°7— 8:2 7:22+-133 0°64 8-86
1g 27 7:°7—18:8 10°85+:-219 1°14 10°50] 26 5:°0— 7:7. 6:67+°149 0°76 11°39
Cc 30 13°8—21-4 18°54+°285 1°56 8-41| 30 4°7— 84 6:°64-°135 0°74 11-14
DM, | 37 *12°2—19°8 “14°7532201 81°77 12:00] 397. «7-2 11-7 «= -: 395-12) ae eg
DM, | 38 18°0—24°:1 20°64+:230 1°42 6:87] 36 11°7—15°7 13°72+°155 0°03 6-77
DM, | 39 28°0—35°9 32:99+°227 1°42 4°30] 37 15°71—18-9 17°14+°171 1°04 6-06
DM? | 36 16-0—20°4 18:37+:240 1:44 7:83] 36 16:3—19°5 17°79+°193 1°16 6°44
DM? | 38 22-2—27-2 24°47+°209 1°29 5°27| 38 19°3—23°1 91-29---169)) 7 -Oo mage
DM? | 38 24:3—29:0 26°33+-191 1°18 4°48] 38 21°-3—26°3 24°:28+-189 1°17 4°82
TABLE 9. Summary of measurements of 331 milk teeth. (N = number of specimens; M = mean
given with standard error; o = standard deviation; V = coefficient of variation).
The breadth/length index has also been calculated (table 10). Here again, the
front teeth and first lower molar show a very high degree of variation, reaching
17°38%. The data indicate that all the milk teeth are longer than broader, the ratio
being close to 100% for the upper molars. They are all relatively longer than the
corresponding permanent teeth (fig. 10), and this feature is responsible for the
lower index (table 10).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 401
DENTAL- INDEX
LENGTH BREADTH B/L INDEX
M D mis: 79.58 Sait: Weil ‘ Sos Tees eaves DENA SUIS TS
M3 py, eee aged ee Be Rete eee ee
Fic. 11. Ranges of variation of the Dental Index for length and breadth and of the
breadth/length index of permanent teeth in Giraffa camelopardalis.
Mean values are indicated.
ANNALS OF THE SOUTH AFRICAN MUSEUM
402
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ASNVUL
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 403
Range of M in corresponding
Tooth N Variation M o Vv permanent tooth
i 22 52°2— 8or1 66-14+ 1-149 6-80 10°28 72-34
ly ag 48°7— 97:0 65°26-+2-187 10°48 16°05 78:69
Ig 26 41°3— 814 62°62+ 1-603 8-16 13°03 71°QI
c. 29 24°2— 44°38 35°78-+ -680 3°66 10°22 41°02
DM, 37 41°Q— QI-2 64°44 1-894. 11°52 17°87 78°59
DM, 36 52°6— 78-2 66-58-+ 1-086 6-52 9°79 Daaly|
DM, 37) 46°:2— 58-0 52°30-+ +509 3°10 5°92 61-21
DM? 35 86-5— 106-6 96-68 -+ 1°035 6-12 6-33 103°QI
DM? 38 78-8— 96-0 86:96-+ -699 4°31 4°95 106-79
DM? 38 82:0—I101°0 92°28-— -741 4°57 4°95 107°44
TaBLeE 10. Breadth/Length (Transverse/A—P) Index of 331 milk teeth, compared with the same
index in the corresponding permanent teeth. (N = number of specimens; M = mean, given with
standard error in milk teeth; o = standard deviation; V = coefficient of variation).
The ‘Dental Index’ has also been determined for the immature teeth (table 11,
He. T 2).
DENTAL INDEX
LENGTH BREADTH B/L INDEX
DMo
DM3
DM3
DM4
Fic. 13. Ranges of variation of the Dental Index for length and breadth and of the
breadth/length index of deciduous teeth in Giraffa camelopardalis.
Mean values are indicated.
4.04. ANNALS OF THE SOUTH AFRICAN MUSEUM
Range of :
Index N Variation © M s Vv
DM? .
Length ue 36 62:7— 81°9 75°38+0-811 4°87 6-46
DM? : .
Breadth == 36 (7551023 83°94+0-678 4°07 4°84
= Transv./A—P DM? F
ndex Transv./A_P DM 306) 94°0— 127°4 110°342-0°629 3°72 3°37
DM?
Length DM: 38 85°7— 102°2 92°93+0°545 3°36 3°61
DM?
Breadth DMA 38 82°4— 92°7 87°47+0°472 2°91 Bae
rade Transv./A-P DMS , 3 ie 3
ndex Transv./A-P DM# 3 51D TOL] 94°11 0°735 4°53 4°01
DM, ae .
Lengtn OM 37 54°6— 96:3 71°05+0:983 5°98 8-41
3
Breadth = 35 56-2— 80-2 68-95-- 0-991 5°86 8°49
3
A Transv./A-P DM, 2 : : an
e€ ———— . —= . . . . °
n ji Transy (AcP ODM, 25 3°7—134°2 90°544 3°113 10°40 i 7
Length awe 38 55°6— 72°3 63:11-+0°581 3°58 5°67
4
DM,
Breadth = 36 7FO-2— O21 79°73+0°821 4°93 6-18
4
Transv./A-P DM,
Index ——————____ 36 98°7—144:°0 123°52+1°760 10°56 8°54
Transv./A-P DM,
TABLE 11. Dental Index for lengths and breadths, as well as the index for the transverse/A—P ratios
of successive immature teeth of Giraffa camelopardalis.
CHAPTER 6
ROOT ANOMALIES
Root anomalies are by no means uncommon among recent giraffes. They will
be dealt with in the adult dentition first, and then in the milk dentition.
I. ADULT DENTITION
Three main types of anomalies have been observed: accessory roots, root
reduction and root fusion.
A. ACCESSORY ROOTS
They are by far the commonest abnormality. Among the 62 adult specimens
analysed, not less than 15 individuals (24%) show distinct accessory roots in a total
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 405 .
of 38 teeth (table 12). The data indicate that the accessory roots are found exclusively
in the last two lower premolars and in the lower molars, the incidence being much
higher in M, and M3.
———$_— |, | ————————_ ———————————————————— —
P; P, M, M, M,
SS ee |
LeR ey ee eens | Deal ped ss L
Xs x De
De x
24291 x x
27137 x x
27752 x x x x
34425 x x
34424 x x x x x
34426 ? ? ? ?
35536 x x x x
51198 x x x x
69403 > 4 x x
83460 x
139696 x x
154033 ?
251797 x
TOTAL 5 =I 4 14 14
e's ff
TABLE 12. Distribution of accessory roots in adults. (R = right,
L = left. In two specimens, it has been impossible to tell from the
records whether the anomaly affects the particular teeth bilaterally or
not: a ‘?’ indicates that the side is unknown).
The most common type is that of a small, thin and short root located between
the two major anterior and posterior roots, either on the buccal or the lingual side.
It never develops the flattened plate shape (side-to-side) of the two main roots. It
is rather circular, hardly extending into the bony socket of the mandible. In some
cases it is poorly developed, resembling a very small tubercle and projecting 2 or
3 mm. from the root base, forming a small depression in the alveolar border of the
mandible. In this series the accessory root is always situated on the lingual side of the
premolars and on the buccal side of the molars. On the premolars it occurs unilater-
ally, whereas—with very few exceptions—the accessory root is bilaterally situated
on the molars when present. In these cases, the right molar presents the anomaly
as the mirror image of the left corresponding tooth.
It is a debatable point whether this root is really an accessory root or whether it
is just the result of some division or splitting off of one of the main roots. On one
specimen only (27137), there isa definite indication of a bilateral division of the
anterior root of M,: on both teeth the anterior root is bifid, the distal branch being -
clearly situated under the protoconid. In all the other cases, however, the accessory
root is central, situated under the protoconid-hypoconid junction, quite separate
from the anterior and posterior main roots.
_ It is not possible to link in any way the molar accessory root with the presence
of an ectostylid: among the 21 cases of molar accessory roots, only 13 correspond to
specimens presenting any form of ectostylid. This does not indicate a significant
406 ANNALS OF THE SOUTH AFRICAN MUSEUM
correlation between accessory root and ectostylid, especially when one considers that
in 86 molars presenting an ectostylid (table 6) there are only 13 which have an
accessory root. ,
B. Root REDUCTION
Root reduction has been observed in three cases only, all P,. In these, the tooth
has only one single root, round in section and rather conical in appearance, instead
of the two normal (anterior and posterior) roots. The specimens are 34423, 34424
and 83460. On the former two, the reduction has only been observed unilaterally,
the corresponding opposite tooth root being perfectly normal. Specimen 83460 has
only one P, (right), the corresponding left premolar being congenitally absent. The
measurement of the teeth concerned in the three specimens are compared with the
mean and range of variation of 109 other P, specimens (table 13):
Transv./A—P
Specimen Taker td Transverse Index
34423 R (reduced root) | 13-0. OG, 88-4
L (normal root) 18°9 15°9 84:0
34424 R (normal root) 14°] 17°5 120°4
L_ (reduced root) 15°5 16-4 105°9
83460 R (reduced root) 13°6 13°5 99°2
L_ (tooth absent) = — —
Series of 109 teeth
Mean 17°8 15°5 87°4
Range of variation 13°3—22°6 8°4—19°4 56-5— 124-0
TABLE 13.
From these figures it may be inferred that, in two of the three observed cases,
reduction of the root corresponds to an appreciable reduction in A—P diameter
(viz. 22°5% and 23:6% in 34423 and 83460 respectively). ‘The reductions in trans-
verse diameter are 21:7% and 12:9% respectively: it is probably not very significant,
especially in view of the negative reduction (—o-5) in 34424. Similarly, in these
three specimens, the A~P measurement is situated at the lower limit of the range of
variation, while the transverse breadth falls within the range of one sigma from the
mean value. Comparing further the teeth with the reduced root with the opposite
ones which have the normal root, there is a distinct correlation between reduction in
root and in size of the crown in one case only (34423). On the other hand, it is
remarkable that, with the exception of the above case, a reduction in size of the
tooth does not seem to correspond to a reduction of the root.
C. RooT FUSION
One single case of root fusion has been noted: the M? in 53546 has (bilaterally)
a bridge completely fusing the posterior buccal root with the lingual one along the
total height of the roots.
2
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 407
II. MILK DENTITION
Root variation is frequently observed in the third lower milk molar DM,. A
total of 20 immature skulls, in which this tooth was still bilaterally in situ and
sufficiently well preserved to allow observations, were studied. All the 40 DM, have
three roots. The two main roots (anterior and posterior) are well developed,
extending bucco-lingually, being rather flattened from side-to-side, and corre-
sponding to the two extreme pillars of the deciduous molar, but they do not show
a very high degree of variability. However, situated between them, there is con-
stantly a third root, corresponding to the intermediate pillar. It is rather buccally
located, circular in shape, and is shorter than the two main roots.
The development of this central root is quite variable. The normal appearance
(so-called because it was observed in 28 out of 40 cases) is that of a cylindrical root,
sufficiently long to penetrate the alveolar socket of the tooth where it is really
‘rooted’, although less than the other two main roots. This is the general rule for
the left DM, (18 cases), and shows in only to of the cases on the right side of the
mandible (table 14).
Variation Right Left Total
‘Normal’ 10 18 28
Reduced single central root I I 2
Divided central root:
2 normal rootlets
I normal and
I truncated
2 truncated
3 truncated
tm NOD iS~)
TOTAL 20 20 40
TABLE 14. Variability in the central root of DM,, and distribution
in right and left teeth.
In one case, this central root was so small and reduced that it hardly indented
the mandible, and could scarcely be called a real root, being not very unlike some of
the accessory roots observed on the permanent molars or premolars (vide supra),
although it was much thinner. In ten cases, however, the central root was bifid or
multiple, possessing two or even three small thin rootlets parallel to each other and
originating from the crown-root junction. Here again there appears to be a high
degree of individual variation, and one could schematically summarize the obser-
vations by distinguishing three main types of ‘multiple’ central root: (i) a double
central root, each of the two parallel rootlets being as long and as large as the normal
single ones: (ii) in addition to the normal central root, and parallel to it, the second
accessory root is much shorter and reduced, hardly reaching the bone where it is
never well ‘rooted’; (iii) two or even three small reduced rootlets are found, truncated
and just reaching or hardly penetrating the alveolar border.
408 ANNALS OF THE SOUTH AFRICAN MUSEUM
CHAPTER 7
APPEARANCE AND VARIATIONS OF THE HORN-CORES
Lydekker (1904) attempts to classify Giraffa camelopardalis, distributed over the
vast geographical area from the Cape to the Sudan and Ethiopia, according to the
colour and the blotching of the skin (see Introduction, page 375) and to the variation
of the horns. He describes a gradual transition from south to north from a two-
horned animal into one (so far as the males are concerned) with three horns, but he
believes that this is by no means regularly progressive, for one finds in the eastern
districts of the Continent a five-horned and even a six-horned ‘race’. On this basis,
Lydekker finds it possible to distinguish:
(a) seven subspecies of the Northern and Eastern giraffes which are all characterized
by the development of a large frontal unpaired horn, namely, typica (in Nubia),
reticulata (in Somaliland), antiquorum (in Kordofan), cottoni (in South Lago,
Uganda), rothschildi (Baringo), tippelskirchi (Kilimanjaro) and congoensis (Congo) ;
(b) four other subspecies of the Western and Southern giraffes, whose frontal horn
is rudimentary or even aborted, namely, peralta (Nigeria), angolensis (Angola),
ward. (northern Transvaal), and capensis (Cape) (fig. 14).
The main paired horns are always present, but particularly well developed in
rothschild: and still better in wardi. Posterior or occipital horns are very well developed
in these two subspecies, but less in cotton. No reliable information on these posterior
formations was available for tippelskircht and angolensis. An unusual ‘azygous’ orbital
horn was described in cotton: and rothschildi, projecting horizontally outwards from
the middle of the frontal border of the right and left orbit respectively. Lydekker’s
observations are summarized in the following table.
Main pair of Additional or
Subspecies Anterior horns horns Posterior horns - ‘azygous’
typica x x — —
antiquorum 54 x — —
reticulata Bre x ? —
cottont x x x x
smaller than in smaller than in on right orbit
rothschildi rothschildt
rothschildi x Ke x sometimes; then
well developed well developed on left orbit
tippelskirchi x x e, —
smaller than in
rothschildi
congoensis a x ? —
angolensis x x ? —
wardi x x x —
large, aborted well developed stronger than in
rothschildi
capensis x x — —
rudimentary
peralta —- x ? —
TABLE 15. Schematization of Lydekker’s subspecific classification and horn distribution.
(x = present; — = absent; ? = doubtful.)
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 4.09
TYPICA
(Nubian desert)
ANT/QUOQRUM
( Sudan )
PERALTA
( Nigeria)
RETICULATA
(Somaliland )
COTTONI \
(Uganda)
ROTHSCHILD!
(Kenya )
TIPPELSKIRCHI
CONGOENSIS (Tanganyika )
( Congo)
ANGOLENSIS
(Angola)
WARD!
(Transvaal)
CAPENSIS
(Cape)
Fic. 14. Map of Africa indicating the distribution of the subspecies of Giraffa camelopardalis,
according to Lydekker’s classification (1904).
Because of the large amount of material available to the authors, the following
observations indicate that Lydekker’s useful information must be somewhat revised.
The sexual difference is very marked in the horns which are usually poorly
developed in females (fig. 15): thus it is hardly possible to make any subspecific
classification on the basis of the horns. The median frontal horn does not exist in
females, there being only a slight swelling on the frontal without development of the
‘ossicone’. The main horns, which are always much smaller than in the males of the
same catalogued subspecies, measure 8-15 cm. in length from tip to base, the dia-
meter not exceeding 3 cm. They are cylindrical and smooth, pointed at their tip, and
410 ANNALS OF THE SOUTH AFRICAN MUSEUM
\
x! x2 20884 34423
eee ey —
53546 53549 82002 83458
83459 162988 163324 270594
Fic. 15. Sketches showing portions of the skulls of female giraffes (Giraffa camelopardalis)
indicating the major variations in the development of the horns. Lateral views,
except X! which is norma verticalis.
usually constricted at their base. Posterior occipital horns and azygous horns were
not found in the females.
ode eee AK ee
TN
A
8377 b 24292 Fy Die we
34422 94425 34426 53550
53765 oe 57675 pais
Fic. 16a. Sketches of portions of skulls of adult male giraffes (Giraffa camelopardalis) showing
major variations in the development of the horns. Views are either norma lateralis
or verticalis.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA AII
Males show a better development of their horns (fig. 16a, b) and it is possible
to notice important differences. But does the wide range of variation permit sub-
specific distinctions? In order to answer this question, each type of horn will be
discussed separately.
Se
81821 81822 82001 8B 83460
/ MY
oa
ea adh 2) '
B 139696 154033 155438 163312
165051 200151 251799 279405
Fic. 16). Further sketches of portions of skulls of adult male giraffes (Giraffa camelopardalis)
showing major variations in the development of the horns. Views are either
norma lateralis or verticalis.
A. SINGLE MEDIAN FRONTAL HORN
It varies from a simple swelling of the frontal (251799) to a true, very strongly
developed horn (155438). The intermediate stages are observed as either a horny
plate (154033), a slight eminence (279405), a globular formation (54123), or a rather
large and regular cone (200151). Usually the anterior part of the horn has a more
gentle slope (155438) which gradually extends on to the nasal bones. Very often on
the median sagittal line, anteriorly, are one (155438), two (81821), three (27752),
four (81820) or more (27137) smaller nodules; one or two may even be found on
the side of the horn (139696) or on the top (81822 and 53765). The profile of the
posterior part of the horn is mostly S-shaped (163312), so emphasizing its knob-like
appearance. The whole formation is generally symmetrical on both sides of the
sagittal plane. In one case however (27137), its transverse horizontal section looked
like a crescent open anteriorly and to the right, with both extremities of the crescent
converging towards a completely isolated smaller knob.
412 ANNALS OF THE SOUTH AFRICAN MUSEUM
B. Parr OF MAIN HORNS
All the male giraffes have these two horns very well developed. The shape is
commonly that of a rounded column, measuring 15-23 cm. in length, 4-6 cm. in
breadth, and the diameter at the tip is 4-7 cm. A difference in size between right
and left horn is by no means rare, the one being sometimes 35% longer than the
other. Both horns are usually parallel: if not, their divergence from the base is
hardly noticeable, so that the external interhorn breadth measured at their tips is
always smaller than the external biorbital breadth. The angle formed by the horns
with the Frankfort plane is very constant and in all the specimens where this could
be measured, it ranged from 48° to 58°. The general shape of the horns is that of a
column regularly cylindrical from the base upwards, and at the tip it forms a rounded
knob (83460 and 24292). The tip is sometimes flattened (27137), the medial edge of
which then projects more than the lateral one. In other cases the column maintains
its regular circumference from base to tip (53550, 1396906).
These fronto-parietal main horns are usually relatively smooth, having small,
shallow vertical grooves, but they may possess a few marked axial ridges (163312
and 8371b), or display small knobs, in series of 2-5, forming irregular crests on the
anterior (57675), medial (27752) or lateral (54123) margin of the horn.
C. PosTERIOR OCCIPITAL HORNS
The posterior ‘horns’ are much more a type of occipital crest or exostosis, con-
densed in two parallel eminences, than real horns. The crest is also very variable
and may be completely absent.
D. SUPPLEMENTARY KNOBS
Smaller isolated supplementary knobs are not infrequently developed either on
the lateral aspect of the frontal, between the posterior supra-orbital border and the
base of the main horns (155438) when there are two or three concentrated in the
same area, or on the orbital border itself (54123) when it is comparable to the azygous
orbital horn described by Lydekker. But in the specimens studied it was distinctly
bilateral and it is a knob rather than a ‘horn projecting outwards horizontally’.
On the basis of the above results, the conclusions of Lydekker relative to the
taxonomical significance of horn-shape and disposition are questionable. It is not
impossible to find typical examples displaying the subspecific characteristics proposed
by Lydekker: male 165051, for instance, from South West Africa, with big main
horns (23 cm. in length) and a very rudimentary ossicone on the frontal. On the
other hand, it is not difficult to find specimens which do not fit in his descriptions,
€.g. 251799, registered as G.c. tippelskircht, which certainly originated from ‘Tan-
ganyika, has no anterior horns and shows just a slight swelling of the frontal.
Similarly specimens 24292, 34422, 34425, 34426 and 83460 which are all males from
Bechuanaland (Mababe Flats) definitely have a very well developed anterior horn
even though they geographically belong to the group of Angola, northern Transvaal
or Cape, which—according to Lydekker’s description—should have no such a large
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 413
horn. Specimens from British East Africa, identified on the basis of their skin as
G.c. rothschildi, have very strong main horns (155438 and 200151) whose length,
general pattern or morphological appearance could not differentiate them from
tippelskirchu. Among both groups one finds long, regularly cylindrical and smooth
horns, or shorter, plumper horns with knobs (163312 and 27752; 54123 and 251799).
Lateral ‘azygous’ knobs are found in both types (27752, 54123 and 155438), but on
the one skull from Uganda, which should be G.c. cottont, no such knobs were found,
as should have been expected from Lydekker’s description.
Consequently, it would be reasonable not to attempt any subspecific determi-
nation of modern African giraffes on the basis of the horn-cores. This principle is
extremely important, as will especially be indicated in the discussion on the horn-
cores of the extinct giraffids.
SECTION II
THE FOSSILIZED GIRAFFIDS
INTRODUCTION
Fossilized giraffid material has been discovered at various sites in Africa,
namely:
Nort ArricA Oran (St. Charles) (Pomel 1892) ;
Bou Hanifia, Chaachas (Reygasse, 1919-20) ;
St. Arnaud (Arambourg, 1934, 1948) ;
Garet (Garaet) Ichkeul (Arambourg, 1949) ;
Douaria (Roman and Solignac, 1934); __
Djebel M’dilla (Arambourg, 1952).
EGYPT Wadi Natrun, Garet-el-Multk (Stromer, 1907);
SUDAN Abu Hugar (Bate, 1951).
East AFricA Omo (Arambourg, 1947);
Serengeti (Dietrich, 1937, 1942);
Olduvai and Kagua (Hopwood, 1934, 1936);
Olorgesailie, Kanam and Rawi (Leakey, 1951).
SouTH AFRicA Vaal River (Haughton, 1922);
Florisbad (Dreyer and Lyle, 1931);
Cornelia and Tierfontein, near Port Allan (van Hoepen, 1932) ;
Makapansgat (Cooke and Wells, 1947; Broom, 1948) ;
Hopefield (Singer, 1954);
some unknown place, probably from the Vaal River gravels
(Cooke, 1949).
Nearly all the above-mentioned material have been either assembled in
Cape Town on loan for study, or examined in the respective Museums where
the specimens are housed. In addition, for comparative purposes, an extensive
survey of the giraffid material from the Siwaliks was carried out in the American
Museum of Natural History (New York) and the British Museum (Natural
History), London. It was not possible to obtain the original specimens
previously described from the following sites outside Africa:
GREECE Veleés (Schlosser, 1921);
Pikermi (Gaudry, 1861, 1867);
Samos, Salonique (i.e. Bounardja and Vateliik) (Arambourg and
Piveteau, 1929).
U.S.S.R. Taraklia (Khomenko, 1913).
Huncary Baltavar (Peth6, 1885);
Polgardi (Kormos, 1911). |
FrANcE Mont Lébéron (Vaucluse) (Gaudry, 1873).
Turkey Adrianople (Abel, 1904);
Maragha (de Mecquenem, 1924).
414
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 415
In respect of the latter group of specimens all the available literature was
studied. All the original material (from Africa) examined will be described
according to geographic distribution.
CHAPTER I
OLDUVAI (OLDOWAY) GORGE, TANGANYIKA
A. GEOLOGY
The history and the geology of the enormous Olduvai Gorge (fig. 17(a),
(b)) have been recorded by Leakey and by Reck, respectively (Leakey, 1951).
Five beds are described: Bed I was deposited at the beginning of the East
African Middle Pleistocene (early Kamasian), Beds II and IV correspond to
the Kamasian and Kanjeran pluvials respectively, the Beds III and V represent
the dry inter-pluvial and the post-Gamblian periods respectively. Giraffid
remains have been recovered from Beds I-IV, although most of the material
comes from Bed II (vide infra).
ATL. ALBERT
5
L.EVASI_? 6
Fic. 17a. Map of Central-East Africa indicating some
of the more important fossil sites.
416
LETTERS REFER TO AREAS OF SITES
NOTE:
ANNALS OF
THE SOUTH AFRICAN MUSEUM
08
HW
PEK
+O
en
“xO
¢
2 s¢
Leen
e
&—
AY
\
MILE
Fic. 175. Map of Olduvai Gorge indicating designations of various sites (from Leakey, 1951).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 417
B. THe MATERIAL
The material recovered consists of loose teeth, or fairly complete dentitions,
postcranial bones (mainly incomplete) and so-called ‘antlers’ (horn-cores).
1. Previously described. Hopwood (1934) described a right fourth lower
premolar (M 14200) as the holotype of Helladotherium olduvaiensis sp. n., and
as paratypes he mentioned a rolled fragment of a right mandibular ramus with
P,-M, (M 14686, horizon unknown) and a partial hind-limb (M 14687,
Bed I). With the discovery of Sivatherium-like ‘palmated antlers’ (inverted
commas, ours) from Bed II Olduvai (M 14954-14955) and from Kagua
(M 14956), Hopwood (1936) referred the above material to Sivatherium oldu-
vaiense. However, Dietrich (1937) stated that on the basis of a metacarpal
(E. 122) which he had studied, he had already referred the Olduvai material
to the Sivatheriinae (although he does not state whether this was ever
published).
2. Observations on material housed in the Coryndon Museum (Nairobt). Material
was kindly sent on loan by Dr. L. S. B. Leakey, Curator of the Coryndon
Museum, and in addition horn-cores and postcranial remains from Olduvai
were also studied in the British Museum (Natural History) and in the Coryndon
Museum (Nairobi). Each specimen is described in detail, the specimen being
numbered according to the museum registers, and the information given below
with each number is found written on each specimen. The inscriptions have
the following interpretation:
Midge :s 2.2: .. OlduvairGorze.
Oldy. .. .. Olduvai (Oldoway) Gorge.
Be =... , ..- Indication.of the site (sec-fig. 17(b)).
Rey ere Soy Bed ir.
1951, 1952, etc. .. Collected during those seasons.
a .. ... The specimen has been found on the surface of the
Bed referred to, but it does not necessarily belong to
that Bed.
M ... .. .. Before a number indicates registration by British
Museum (Nat. Hist.).
Marsabit Road .. A site about 100 miles east of the southern point of
Lake Rudolph in Kenya.
Coryndon Museum Guraffid Material
A. Horn-cores
86 ns .. Old. BK II 1952.
Old. 1.53 .. (Provisional number given by Mrs. Coryndon, in 1956.)
Old; ot5s\ 2. (ddem.)
Old. 1952 .. SHK II BK II base (S).
M 149545 .. Oldy. BK IIS (13.V.935).
Old. 3.53... (Provisional number given by Mrs. Coryndon in 1956.)
M 17027. .. Oldy. BK IIS (14.V.95).
M 17028 .. Oldy. BK IIS (14.V.35).
M 17029.) «. Oldy. BK ITS (14.V.935).
M 14955 ~.. + (At present in British Museum (Nat. Hist.).)
B. Fragments of jaws
Upper:
F 3655 .. Right maxilla, M!—M4&, Old. II, 1941, in situ.
ANNALS OF THE SOUTH AFRICAN MUSEUM
Lower
6 .. .. Right mandible, P,—M;, Old. BK II.
F 3656 .. Left mandible, M,—Ms, Old. II.
I .. .. Right mandible, P,—M,, Old. BK II.
365.. .. Right mandible, M,—P;, Old. SKH II, 1953.
g92.. .. Right mandible, M., Old. SHK II, 1953.
3 0.. “2 Right mandible M5. Old2BK IL
93 .. .. Left mandible, M,—M,, Old. BK II, 1952.
g2.. .. Left mandible, M,—Ms3, Old. BK II, 1952.
gI . Left mandible, M,—M,, Old. BK II, 1952.
Neguntiri site. Left mandible, M,— Mg, Old. BK II East 1953.
g21.. .. Left mandible, M,—M,, Old. BK II, 1953.
C. Isolated teeth
Upper:
premolars.. F 2989 ?P%, Oldoway 1941 II, in
molars .. F 2993 M?, Old. II.
4 M$, Old. BK II.
109)=—s- M, Old. BK II, 1955
F 2992 ?, Old. II in situ.
Lower:
canine, 2. 97 Old. BK II, 1952.
premolars... F 2991 P,, Old. IS, 1941.
2 P,, Old. BK II.
5 P,, Old. BK IL;
7 Pi Old BK AL.
96 P,, Old. BK II, 1952.
molars .. 105 M,, Old. BK II, 1953.
132 M,, Old. BK IT, 1953.
95 Mz, Old. BK II, 1952.
120 Ms, Old. BK II, 1955
— M,, Marsabit Road.
Fragments of molars, further unidentifiable
166 Old. BK II, 1955.
116 Old. BK II, 1953.
8 Old.
too )=6©- Old. BK _IIT,, 1953.
g8-—99 Old. BK IT, 1952.
= Marsabit Road.
— Olduvai surface.
D. Post-cranial skeletal remains
Forelimb:
116p.. .. Proximal end of ulna. Old. BK II, 1952.
115.2 ..».) Distal end of radius? Old. BK II, 1952.
941 =... ~ Os magnum. Old. BK IT, 1952.
114... .. Distal end of metacarpal. Old. BK II, 1952.
Hindlimb:
100" )..5.. - Distal end of feniur- Old. BK IT, 1952.
101, 112 .. Distal end of tibia. Old. BK IT, 1952.
103, 108 .. Calcaneum. Old. BK IT, 1952.
102,107 .. Astragalus. Old. BK IT, 1952.
104, 109, 110 Cubonaviculare. Old. BK II, 1952.
105, 110A .. Cuneiform. Old. BK II, 1952.
106, 111 .. Proximal end of metatarsal. Old. BK II, 1952.
Blanes .. Distal end of metatarsal. Old. BK II, 1953.
M 14687 .. Almost complete articulated hindlimb, in B. M. (N.H.).
Proximal phalanges:
113 .. Old. BK II, 1952.
185 »»,, Old. BR IT. 1953;
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 419
B:g097 :... Old. SII.
Peob4. s.. Old. Sif, 1948.
E965) J.
ew, ic» Old. surface.
Sesamoid bone:
gaa Sh.” ‘Old’ TI, 1952.
DESCRIPTION OF SPECIMENS
A. Horn-CoRES
86. Old. BK II 1952 (plate 1(d), (f)).
This is a right horn-core. The base is hollow, extending about 170 mm.
in depth from the broken edge. In section it is roughly triangular, the rounded
apex being posterior. This posterior border becomes increasingly rounded
towards the tip, while the more anterior region, rounded at the base, narrows
to a well-defined border towards the apex. The anterior surface has knobs on it.
The medial surface is convex, both transversely and medio-distally, and
presents deep grooves. Three of these are parallel to each other and to the
convex anterior border, along which they run, even on to the knobs, following
their profiles. Two other grooves are some distance from the anterior border,
passing parallel to each other upwards and towards the posterior border.
There is also a short, very broad, oblique groove which leaves the base
at the anterior convex border and runs obliquely upwards and backwards on
the medial aspect.
The general curvature and the rotation (twist or torsion) of the horn-core
are not marked: it is almost straight.
No flange* is visible. This portion has been reconstructed in plaster but
it would appear unlikely that a flange would have been present. The first
visible knob has been called knob 2.
This specimen is very similar to the horn-core from Garet Ichkeul (see
Section ITI, chapter 1).
Length along posterior (concave) border: 710
Length along anterior (convex) border: 810
Height of the knobs: 20-40
Circumference A-P Breadth*
At base 415 150 114
Just under knob 2 350 128
At knob 2 145 89
Just under knob 3 330 11g
At knob 3 130 83
Just under knob 4 300 113
At knob 4 135 78
Just under knob 5 280 106
At knob 5 110 64
At ‘tip’ (broken) 75 63
(* Side-to-side diameter.)
TABLE 16. Measurements of Old. 86 (mm.).
: * The term ‘flange’ is applied to the appearance often presented by the anterior border
just above the base when it narrows, flattens and sweeps (flanges) outwards.
420 ANNALS OF THE SOUTH AFRICAN MUSEUM
Old. 1.53 and 2.53 (plate 1(c), (e); 1 (a), (d)).
1.53 1s a right horn-core and 2.53 a left. They are very similar to one
another, although they may not belong to the same individual because the
rotation (torsion) of the horns differ from each other, 1.53 rotating go° anti-
clockwise, while 2.53 rotates clockwise (normal) through 140° from base to
tip. This indicates the tremendous variation in torsion between horns which
are otherwise very similar. Both horns have a marked flange (267 mm. and
155 mm. in length respectively) which terminates superiorly in knob 2 above
which there are three other knobs (3-5) along the antero-medial border. The
flange of 2.53 is shorter than that of 1.53. On the flange of 1.53 there are four
small knobs, the highest of which corresponds to knob 2. These are not present
on the flange of 2.53. On both horns there is a knob opposite the middle of the
flange, but it is situated on the posterior border somewhat medially. This knob
is designated ‘P’ and may be compared with the blade-like projection which
led Falconer (1868) and Abel (1904) to term the horn a tri-radiate
antler.
In both specimens the anterior border first passes upwards and laterally
from the base (where it is ill-defined) and at about the middle of the flange it
swings towards the medial side. In this way the lower part of the horn develops
its rotary twist, which is more obvious in 2.53. Thus the anterior border which at
the base is really antero-lateral tends to twist to the antero-superior aspect of
the horn, while the posterior border—which is postero-medial at the base—
tends to become inferior at the tip. Furthermore, as a consequence of this
torsion, the antero-medial surface (which has the marked grooves) comes to
lie on the posterior aspect of the horn above the flange region.
In specimen 1.53 the postero-lateral surface has no grooves at all, while
the antero-medial surface presents two grooves near the anterior border, both
parallel to it, and both broad but not deep. The cranial fragment at the base
of 1.53 contains a few shallow sinuses but does not display any sutures and it is
not possible to assess the correct orientation of the horn to the skull. It would
have been valuable to obtain an understanding of the position of the horn on
the skull so as to determine whether the criteria the authors have utilized for
‘siding’ the horns are correct or not (see also Section III, chapter 1).*
The base of 2.53 is hollowed out to a depth of approximately 300 mm.
from the broken edge. There are some sinuses present. The grooves resemble
those of 1.53 but they are more numerous, there being four on this specimen.
Distances between the knobs 1253 2.53
2—3 141 196
54 110 164.
4—5 Uc k4o 146
* At the galley-proof stage, Dr. L. S. B. Leakey reported to one of the authors (R. S).
that a Sivatherine skull complete with attached horn-cores had just been discovered at Olduvai
and that a note on it would soon be published by him.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 421
Length of the flange 267 155
Length along anterior (convex) border 950 IOIO
Length along posterior (concave) border 840* 690
* A portion of skull attached to the horn is included in this measurement.
Circumference A—P Breadth*
I-53 2293 £53 2-53 T5535: 2°53
At base 400 390 159 145 127 cere)
Middle of flange 410 420 154 172 79 81
At knob 2 380 410 145 165 84. 89
Just under knob 3 300 290 105 IOI
At knob 3 330 290 129 104. 76 75
Just under knob 4 280 250 106 go
At knob 4 290 260 rie: 97 67 63
Just under knob 5 270 250 99 97
At knob 5 290 290 bry 115 59 58
Tip 170 170 52 59 52 47
(* Side-to-side.)
TABLE 17. Measurements of 1°53 and 2°53 (mm.).
Old. 3.53 (plate 2)
This is a fragment of a left horn-core with part of the proximal portion
missing. The tip and distal end are practically intact. At the base the flange is
still present, ending in knob 2. Above this is a large knob 3, the only other knob
on the anterior border.
The hollowed-out portion of the base is very narrow and extends about
130 mm. from the broken edge. On the antero-medial surface there are two
deep grooves parallel to the profile of the knobs. There are also two more
regular parallel grooves near the posterior concave border. They all extend
to the tip. In addition, there are a number of small grooves at the proximal
end which peter out in the region of the upper end of the flange. On the
lateral surface there are some broad, but less marked, grooves. As in most of
the other specimens (except that from Tierfontein) the edges of the grooves
are irregular and heaped-up in parts.
The fragment, in general, is very similar to the corresponding portion of
the Tierfontein horn-core (C 431). The curvature and rotation are rather
poorly expressed. The region above knob 3 is almost straight.
Length along anterior (convex) border 700+
Length along posterior (concave) border 430-+
Circumference A—P Breadth*
At base 360 138 —
Middle of flange 360 146 77
At knob 2 380 155 73
Just under knob 3 310 122 70
At knob 3 360 14! 68
100 mm. above knob 3 250 89 67
At tip 150 52 43
(*Side-to-side.)
TaBLE 18. Measurements of 3:53 (mm.).
422 ANNALS OF THE SOUTH AFRICAN MUSEUM |
Old. 1952 SHK II BK II base (S) and M 14954b Oldy. BK IIS. (Plate 3(c), (d))
These are two closely fitting fragments belonging to the same left horn-
core. M 14954b noted by Hopwood (1936), but he did not include the ‘b’.
The flange exhibits a small knob 1 at its centre along the anterior border,
Beyond knob 2 (at the upper end of the flange) are knobs 3, 4, 5 and 6.
Distances between the knobs (mm.)
I j=- (2: 83
2 taceral io 185
3 — 4: 140
4 — 5 Ta 1
5 — 6: Sel
6 — tip: 174
There are a number of nutrient foramina (as in 1.53 and 2.53) which are
not very large (2-3 mm. in diameter) and not specifically related to the grooves.
It exhibits the same type of curvature and torsion as 1.53 and 2.53 but
they are much more twisted and curved. In turn, it shows more torsion than
Old. 86 and 3.53. Thus it occupies an intermediate position and this series
indicates a further grade of variation of the torsion of the Sivatherine horns.
As in 1.53 and 2.53, there is a medio-posterior knob (‘P’) on the inferior
portion near the base and opposite the flange.
The grooves on the antero-medial surface are large, similar to those in
the Hopefield specimens.
The base contains a few sinuses and is hollowed out to a depth of 80 mm.
The horn closely resembles M 14955 (vide infra).
Length along anterior (convex) border 840
Length along posterior (concave) border 580
Circumference A—P Breadth*
At base 330 117 100
Flange (knob 2) 350 143 78
At knob 3 360 14! 66
Just under knob 4 270 102 aa
At knob 4 290 112 62
Just under knob 5 240 go _
At knob 5 260 94 64
Just under knob 6 220 80 ~
At knob 6 240 85 59
Tip 150 49 46
(*Side-to-side. )
TABLE 19. Measurements of M 14954 (mm.).
M 17027 Oldy. BK IIS.
The tip and distal portion of a right horn-core. The only interesting
features are that the extreme tip is somewhat recurved posteriorly and that the
grooves on the antero-medial surface have extended to the base of this recurved
portion where they end abruptly. The extreme tip is rather rough and
irregularly rounded.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 423
Measurements of M 17027 (mm.)
Length of fragment 164
Circumference A—P Breadth*
Broken end 180 63 59
Tip — 36 36
( * Side-to-side.)
M 17028
A fragment of bone. It is difficult to be certain whether it is of a horn-core
or not. The inner, smooth surface may represent a portion of the endocranial
occipital region.
M. 17029 Oldy. BK IIS.
This is the anterior border of the flange with two knobs. There is also a
smaller knob (P) opposite the flange. A fairly deep groove is visible near the
knobs.
Measurements of M 17029 (mm.)
Distance between knobs 1-2 95
Total length of fragment 228
Breadth of flange (side-to-side) 51
M. 14955 (housed in the British Museum (Natural History) (plate 3(a), (0))
This is a left horn-core, the knobs pointing antero-laterally and the
grooves being on the antero-medial surface. Part of the base is broken away,
so that the central (hollowed) part formed by the cranial sinuses is not dis-
tinguishable. The posterior portion of the base is rounded, while the anterior
flange portion narrows to a ridge. Just above the lower end of the horn-core
there is a broad flattened flange, the upper portion of which is broken away in
a V-shaped notch just below a knob-like projection on the antero-lateral border.
This flange has an S-shaped, curved anterior border which arches forwards,
upwards and laterally; then upwards, medially and forwards; and then
upwards, medially and backwards. The antero-lateral border of the horn also
has two other knobs, and there is a small rough projection just above the
highest knob. Just above knob 2 the horn becomes more circular and tends to
narrow (A-—P) rapidly. The tip of the horn is broken off; but it appears to have
pointed horizontally, medially and backwards.
This specimen is noted by Hopwood (1936).
There are deep grooves on the antero-medial surface, which sweep up
from the base more or less parallel to the anterior and posterior borders. On the
postero-medial surface of the horn-core there are less marked grooves present,
some of them being along the axis of the horn, while others are transverse.
4.24. ANNALS OF THE SOUTH AFRICAN MUSEUM
Length along the posterior border .. ENS 2¢ (4 568
A-P length across the broadest part of the flange Bye nt LGO
A-P length at the highest part of the flange (knob 2) oh OE
Breadth (side-to-side) at highest eae of paeee ee Bi ay 85
A-P length at knob 3 te i
Breadth (side-to- cg} at knob 3 ae oe ie uy, 54.
A-P at knob 4 . Aye Ne ia She 88
Breadth (side-to- side) at knob 4 a te his 23 53
A-P at distal extremity of the horn .. oo 46
Breadth (side-to-side) at distal extremity of the horn s 40
TABLE 20. Measurements of M 14955 (mm.).
B. FRAGMENTS OF JAWS
F 3655 Oldoway II: in situ 1941, with others marked ‘A’ (plate 4)
It consists of a large fragment of a right maxilla, a portion of the right
palatine bone also being intact. It contains M3, M? and M!. The teeth are
in a very worn state; M! in particular has been worn almost to the crown-root
junction on its anterior pillar.
Buccal surface. ‘The cingulum is rather marked especially in M*. In M?,
however, the buccal surface is almost completely broken away. Continuous
with the rounded cingulum in M? and Mz? is a very prominent parastyle, a
mesostyle and a metastyle, the parastyle being particularly large and the
metastyle being rather rounded. ‘The metastyle of M? is almost absent, being
heavily impacted against the anterior pillar of M%, and the enamel in that
region is also worn away. The prominence of these styles is most marked on
the third molar.
Lingual surface. Rather coarse rugosity. On each tooth there is a cingulum
below which there is a slight bulge.
Occlusal surface. ‘The central pits of M? and M? are U-shaped, although
the enamel on the buccal aspect of the pit tends to be rather V-shaped. In M?
the enamel of the central pit of the posterior pillar is irregular on the lingual
side. In M?, where there is more wear, the limbs of the pits which sweep towards
the styles are more or less obliterated; and this is even more obvious in M}!
where the central pit is completely obliterated in the anterior pillar, while the
posterior pillar has a small irregular island of enamel remaining, representing
the central pit.
On the posterior surface of the posterior pillar of M3, there is a sharp
indentation of the enamel which is also reflected on the root. The posterior
buccal root of M? is massive and triangular. The buccal roots of the teeth can
be seen as the bone is broken away over them, while only the base of the lingual
root can be seen, and there are no significant differences in appearances
between these and the usual giraffid pattern (vide supra).
6 Old. BK II (plate 7).
The body of a right mandible, containing M. g-P,. The vertical axes of
M, and M, are tilted forwards. ‘The enamel of all the teeth is fairly rugose
(except for P,).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 425
M, has a rounded cingulum. The rugosity cannot be determined as the
enamel is covered by a thin layer of breccia. There is a rudimentary ectostylid
and protostylid similar to those of Old. 1 (vide infra). The talonid is slightly
angulated to the A—P axis of the pillars in a buccal direction.
M,, M, and P, have features similar to those of Mg.
P, is similar to P, of specimen 365 (vide infra), except that the paraconid
is more separated from the parastylid by a ‘pinching in’ of the enamel.
P,
Buccal surface. Finely rugose. There is a rounded cingulum.
Lingual surface. There is a slight cingulum. The parastylid and the meta-
stylid are present.
Occlusal surface. The tooth-shape is triangular. The buccal surface is
slightly rounded and the apex of the tooth points anteriorly. The posterior
surface of the tooth is markedly worn away.
Wear. ‘The teeth are in advanced wear. On the anterior surface of Ms,
the enamel is thinned by ‘impaction’ of the posterior surface of M,. On the
anterior surface of M, the enamel is almost completely worn away; a similar
thinning is seen on the contiguous surfaces of M, and P, where the enamel is
completely absent. The contiguous surfaces of P, and P, show loss of enamel,
as does the posterior surface of P,, but the anterior surface of P, has its enamel
only slightly thinned. Maximum wear on the premolar is found on the posterior
occlusal surface on the buccal side.
Roots. ‘Those of P, are broken away on the buccal side, but there are two
roots rounded in shape and they are continuous with each other on the area
visible above the alveolus, which is about 10 mm.
F 3656 Oldoway 1941, S II (plates 5(b), (d); 6(a))
A portion of a fragmented left mandible, containing M,, M, and a part
of M,.
M; |
Buccal surface. There is a cingulum on the posterior pillar and talonid, but
on the anterior pillar there is a slight depression in the cingulum region and there
is an unusual ridge of enamel a few millimetres above the crown-root junction.
The rugosity is coarse. The tooth is in a rather advanced stage of wear.
Lingual surface. Rather rugose. Marked cingulum formation. On the
posterior surface of the talonid the ‘abnormal cingulum’ arrangement is also
present. Leading up from the cingulum there is a slight entostylid and a
slightly more marked parastylid, while the metastylid cannot be detected.
The median costa of the entoconid is more obvious than that of the metaconid.
The talonid is rounded and its axis is deviated buccally at an angle of about
40° to the A-P axis of the two pillars.
Occlusal surface. ‘The two sides of the central pit of the anterior pillar are in
close contact, except anteriorly where they are separated by a small triangular
426 ANNALS OF THE SOUTH AFRICAN MUSEUM
space. The sides of the central pit of the posterior pillar are slightly more
separated than in the anterior pillar, and there is a protrusion of enamel in the
direction of the talonid. ‘The anterior pillar is more rounded than the posterior
pillar, the peripheral enamel of the posterior pillar being more V-shaped than that
of the anterior pillar, which is rather U-shaped. Enamel of the anterior surface,
where in contact with the posterior pillar of Mg, is almost completely worn away.
M,
Buccal surface. ‘There is a cingulum with a rounded bulge above it. Coarse
rugosity. Its characteristics are generally similar to those of M;. The posterior
surface of M, is worn away where it meets M,, and the anterior surface where it
meets M, is very worn, as is the posterior surface of M,, at least half of the
surface of which has no enamel whatsoever.
M,. A part of the anterior pillar is missing.
1 Old. BK II (plate 6(c), (d), (e))
This specimen consists of a portion of the body of a right mandible con-
taining M,—P, in a fairly advanced stage of wear.
M;
Buccal surface. ‘The enamel is coarsely rugose. There is a rounded cingulum
which is particularly marked ‘on the anterior pillar and the talonid. There is
a very large ectostylid; and there is a slight bulge above the cingulum of the
anterior pillar, but the posterior pillar does not show this and near the occlusal
surface of both there is a slight concavity of the enamel surface.
Lingual surface. ‘The enamel is coarsely rugose. The cingulum is present,
forming a rounded bulge in the region of the base of the entoconid. ‘The
features are similar to M, of Old. 6. The lingual surface is cracked and the
whole crown has been forced lingually.
Occlusal surface. No additional features, except that the enamel is very thick.
M,. On the buccal surface the features are similar to those of M;. On
the lingual surface the enamel of the posterior pillar is broken away. The
appearance otherwise is similar to that of any M, described here.
M,
Buccal surface. There is a great similarity to M,, except that the bulge
above the cingulum is more rounded. A distinct nodular ectostylid is present.
The other aspects of the tooth are similar to those of any M, described here.
P,. The general description is identical to that of P, of specimen 365.
365 Old. SHK II, 1953 (plate 8)
Fragment of a right mandible containing M,, P, and P,;. The teeth are
in a very advanced stage of wear.
M,
Buccal surface. ‘The enamel is rather rugose, and the cingulum is present.
On the posterior surface of M, all the enamel has been worn away, the contact
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 427
surface with M, being formed by dentine, which itself has been worn away,
forming a notch in this surface.
Lingual surface. The enamel has a rather rugose appearance. There is a
small cingulum present. No features can be identified because of the marked
wear and fragmentation.
Occlusal surface. ‘The central pits are completely worn away, except for a
small island of enamel in the posterior pillar. ‘The buccal enamel of the posterior
pillar is U-shaped, while that of the anterior pillar is V-shaped.
Py
Buccal surface. Rather rugose enamel. A cingulum is present, being
particularly marked on the hypoconid. A small ectostylid is present between
the hypoconid and the protoconid.
Lingual surface. Rather rugose. Cingulum present. The base of the
entostylid can be seen. There is a rounded bulge forming the base of the
metaconid. The enamel of the posterior surface is worn thin, while that of the
anterior surface is worn away completely.
Occlusal surface. ‘The central pit of the anterior pillar is V-shaped, while
between the anterior and the posterior pillars there is a small triangular pit.
The buccal enamel of the anterior pillar is convex and wide, “boat-shaped’,
while the buccal enamel of the posterior pillar is typically in the form of a ‘U’
compressed from front to back.
Roots. On the lingual aspect, the bases of the roots can be seen. Thus
two triangular roots are visible, the anterior being slightly larger than the
posterior and they are separated by an inverted V-shaped interval.
Ps
Buccal surface. Coarsely rugose. The cingulum is present, being particularly
rounded in the region of the hypoconid.
Lingual surface. Rather rugose. There is a marked rounded cingulum,
being particularly marked at the base of the entostylid and not quite as marked
at the base of the parastylid. There is a distinct bulge in the region leading
up to the metaconid.
Occlusal surface. ‘The whole tooth gives the appearance of an irregular
triangle, the buccal surface being more convex than the lingual surface, the
apex pointing anteriorly. The posterior surface has its enamel worn away
completely, and the dentine has been hollowed out by P,. The posterior ‘pillar’
has an irregularly shaped triangular central pit, while all that remains of the
central pit of the anterior ‘pillar’ is a small circular island of enamel surrounding
a small pit at the posterior end.
Roots. The root of the anterior ‘pillar’ has a broad triangular base. It is
larger than the root of the posterior ‘pillar’ which is separated from it by an
interval shaped like an inverted V.
Mandible. Because it is highly fragmented and almost completely filled with
plaster, it is considered advisable not to take any measurements of the mandible.
428 ANNALS OF THE SOUTH AFRICAN MUSEUM
392 Oldoway SHK II E, 1953 (plate 9)
Fragment of a right mandible containing M, and the sockets of M, and M,.
M,. In an advanced stage of wear.
Buccal surface. ‘The enamel is rather rugose. The cingulum is present, and
there is a rounded bulge above it.
Lingual surface. Rugosity is not clear because the tooth is worn smooth on
this surface. The cingulum is present. The entostylid and parastylid are
present and are continuous with the cingulum at their bases. A metastylid is
not present.
Occlusal surface. ‘The lips of the central pit are more widely separated in
the posterior pillar than in the anterior.
Roots. ‘The shallow sockets indicate that the roots were short.
3 Old. BK II (plate 10)
Fragment of a right mandible with M;. The tooth is in a fairly advanced
stage of wear.
Buccal surface. ‘There is a marked rounded cingulum. Enamel rather
rugose. Hypoconulid and ectostylid are present as small nodules. The enamel
of the anterior surface of the tooth is broken off, as also the enamel of the
lingual surface of the anterior and posterior pillars. The talonid is rather
rounded and its axis is in direct line with the longitudinal axis of the anterior
and posterior pillars. ‘There is quite a marked V-shaped groove between the
enamel surface of the pillars, and also between the posterior pillar and the
talonid.
Lingual surface. The enamel of the talonid is rather rugose, and if the
talonid is looked at from the posterior aspect the enamel can be seen to bulge
and drape down on the buccal aspect in the typical ‘apron’ effect. The lingual
surface of the talonid slopes rather markedly buccalwards and upwards towards
the apex from the bulge above the cingulum. The buccal surface has a con-
cavity just above the rounded cingulum.
Occlusal surface. The central pit of the anterior pillar is irregularly U-
shaped and closed, while the central pit of the posterior pillar is closed off
anteriorly by the enamel of the posterior surface of the anterior pillar, and
posteriorly the pit is open and continuous with the central pit of the talonid.
Mandible. Breadth opposite talonid: c. 42 mm.
93 Old. BK II, 1952 (plates 5(a), (c); 6(d))
Portion of the left mandible, containing M, and M, and a piece of M,.
The teeth are in a most advanced stage of wear, and the wear is extremely
irregular in that the lingual cones of the pillars of M, and M, have been com-
pletely worn down beyond the crown-root junction, while on the buccal aspect
a fair amount of enamel is still present. Because this type of wear is the reverse
of the usual, it is probable that the cause is a pathological one.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 429
VE
Buccal surface. The enamel is rather rugose, although it is worn smooth in
parts. There is a bulge above the cingulum on the pillars and on the talonid.
Lingual surface. Only the enamel of the talonid and that part where it
communicates with the posterior pillar is visible. The talonid is placed at a
peculiar angle to the rest of the tooth. Not only is its A—P axis angulated
buccally in relation to the mesio-distal axis of the anterior and posterior pillars,
but it is also tilted upwards so that its occlusal plane is angulated at 20° to the
occlusal plane of the pillars.
Occlusal surface. The central pits of the anterior and posterior pillars are
still visible.
M,
Buccal surface. ‘The buccal portion of the posterior pillar and the roots are
broken away, and the surface of the anterior pillar has a rather rugose
appearance and a small cingulum.
Lingual surface. The shape of the tooth has been completely disfigured by
the abnormal mode of wear, as in the case of M3.
Occlusal surface. There is a small island of enamel projecting above the
surface in the region of the parastylid. |
M,. A portion of the buccal surface of the posterior pillar remains, this
pillar being in extreme wear. The rest of the tooth is fragmented and broken away.
92 Oldoway, 1952, BK II (plate 11)
A fragment of a left mandible, containing M, and M, and a piece of the
posterior root of M,. The teeth are in early wear and the crown-root junction
has not yet appeared above the alveolar surface.
M;
Buccal surface. ‘This surface is coarsely rugose. The median part of the
hypoconid and of the paraconid is rather angulated, especially near the
occlusal surface.
Lingual surface. A portion of the mandible is broken away uncovering the
crown-root junction where a cingulum can be seen. Just above it there is a
bulge, and leading up from the cingulum on the anterior pillar there is a
marked narrow parastylid, which does not quite reach the occlusal surface of
the tooth. About half-way up from the crown-root junction, the base of the
-metastylid commences and near the occlusal surface it becomes a prominent
ridge overlapping the anterior portion of the entoconid. The median ridge of
the metaconid is prominent for about the same distance as the metastylid. The
median ridge of the entoconid is prominent, but the entostylid is much less
marked. The lower % of the lingual surface of the enamel of the anterior and
posterior pillars are fused in the region of the metastylid, but the upper 4 is
separated by a groove. This surface of the entostylid is continuous with that of
the talonid.
430 ANNALS OF THE SOUTH AFRICAN MUSEUM
Occlusal surface. ‘The tooth is in early wear which is most marked along
the anterior portion of the protoconid. The central pits are V-shaped, their
hollowed portions being continuous with each other in the region between the
two pillars, while the central pit of the posterior pillar is continuous with a
small central pit of the talonid. The central pit of the anterior pillar is closed
off anteriorly where the enamel of the paraconid fuses with that of the proto-
conid.
M,. The general characters of M, are similar to those of Ms, except that
the entostylid and parastylid are more marked than in M,, and the metastylid
presents a vertical groove which kinks it posteriorly.
g1 Old. BK II, 1952
This specimen consists of the posterior portion of the left body of a mandible
including the angle, and contains M,, a part of M,, and the roots of M3.
Owing to previous inaccurate reconstruction M, appears to be lower than M,.
M,
Buccal surface. Coarsely rugose. Cingulum present. The enamel is
thickened above the cingulum of the posterior pillar. A small ectostylid is
present. The tooth is in a medium stage of wear.
Lingual surface. Finely rugose; cingulum present. It has the general
characters of M, described previously. A portion of the metaconid is absent.
Occlusal surface. Shows two central pits; their lips are fairly widely
separated, the anterior pit broadening out anteriorly and the posterior pit
broadening out posteriorly. The enamel of the anterior surface of the anterior
pillar is worn rather thin due to contact pressure of M,.
M,. The general description is similar to that of M,, but it is markedly
impacted against M,, so that its entostylid appears rather squashed.
Oldoway BK II—East Neguntini site, 1953 (plate 12)
Fragment of an unnumbered left mandible containing M,, M, and the
anterior pillar of M,. It is in an advanced stage of wear.
M;
Buccal surface. There is a small rounded cingulum above which the buccal
surface is flat. Fairly rugose.
Lingual surface. 'The enamel is chipped away.
Occlusal surface. The anterior pillar has a rounded appearance. The
parastylid is small.
M,
Buccal surface. Rounded cingulum above which the enamel bulges fairly
markedly forming a ridge. Fairly rugose enamel.
Lingual surface. It has the general characters of other M, described
previously.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 431
- Occlusal surface. ‘The central pits of M, are linear. The posterior pit has an
extension up towards the entostylid.
M,. Rather similar in general appearance to M,.
321 Old. BK II Ex. 1953 (plate 13)
Fragment of a left mandible containing a portion of an anterior pillar of
M, and a fragmented M,. It is at the same stage of wear as the Nguntiri
specimen.
M,. The buccal surface of the anterior pillar is rather flattened out and
rounded. It has a finely rugose enamel.
M,. It is in rather advanced wear.
Buccal surface. ‘There is a sma]l cingulum which has a small rounded bulge
above it on the anterior pillar. Finely rugose enamel. The anterior surface of
the anterior pillar is broken away.
Lingual surface. Fairly rugose. Cingulum present leading up to reach the
parastylid and entostylid. The parastylid is prominent.
Occlusal surface. It presents an identical appearance to any other molar
at a similar stage of wear. The enamel of the posterior surface of the posterior
pillar is slightly thinned out due to contact wear by M3.
C. IsoLATED TEETH
F 2989 Oldoway, 1941, II, in situ with ‘A’ (plates 15(e); 16(e); 17(e))
This is an isolated upper left premolar, probably P® (vide infra), in an
extremely advanced stage of wear which is reminiscent of that of M! of F 3655.
The lingual surface is completely worn away, while the buccal surface presents
a fairly rugose enamel. Just as the wear is more marked on the lingual surface,
so it is more marked on the anterior surface. The base of a rather large and
rounded metastyle is present.
Occlusal surface. Only a small slit remains of the central pit, as well as a
small island of enamel near the metastyle.
Roots. ‘The lingual root, which is broken off near its tip, is enormous with
a rather convex lingual surface and a flattened buccal surface. There are two
buccal roots, the anterior one being oval, the posterior one being triangular.
The anterior root is broken off near its tip.
The measurement of the mesio-distal length of the root, just above the
crown-root junction is 27-1 mm. which compares favourably with the length of
the root of the Hopefield P* (4025), viz. 29°7 mm.
Determination of the diagnosis of F 2989
In order to determine which upper premolar it is, the premolar index
(see Section I) has been calculated.
Morphologically, specimen F 2989 seems to belong to the same individual
maxilla as F 3655. Both were found in 1941, ‘with A’, and they look very
432 ANNALS OF THE SOUTH AFRICAN MUSEUM
much the same, i.e. degree of wear, type of fossilization, colour of tooth and
breccia.
43
a)
The premolar index for the transverse breadth is — — one
47
In G. camelopardalis, the same premolar index for the transverse breadth is
Range of
N IMD (Sc sce; G Vv Variation
P3
aE 108 91:03-+ -469 4°84 5°31 73°6—106-2
pt
a 108 95°74 458 4°72 4°93 83°0— 109°5
- According to these figures, the premolar index of F 2989 is more closely
related to that of P?/M? than to that of P4/M!, and it should rather be considered
asayb?:
However, it should be noted that 91-48 is different from the mean value
of P4/M! by less than one sigma (95:74—4:72 = 91:02) so that no conclusive
significance could be attached to this mathematical approach of the problem
of determination, especially in view of the large range of variation.
F 2993 Oldoway II, with ‘A’ (plate 14(a), (0d), (c))
Isolated upper right molar, probably M?. It is in a fairly advanced stage
of wear, and has a rather rugose enamel, and a cingulum which is very marked
on the anterior pillar. There is a small entostyle and hypostyle.
Buccal surface. The paracone is missing, but the metacone shows the
usual characters except that the cingulum is a very marked ridge, and that the
central costa of the metacone is not quite confluent with the cingulum at its
base. There is a heaped-up ridge of enamel just above the cingulum.
Lingual surface. Below the rolled lower edge of the cingulum, the enamel
appears to be thrown into folds.
Occlusal surface. It is typical of the upper molars with no unusual features,
except that the enamel surface of the hypocone is quite widely separated from
that of the protocone where they tend to come together between the two pillars.
4 BK II (plates 15(b); 16; 17)
Isolated right M%, with the metacone broken away. It is in a most
advanced stage of wear.
Buccal surface. There is a marked cingulum. The typical formation of the
styles are seen here again, but the ridge of enamel leading to the paracone is
not very marked.
Lingual surface. The enamel is fairly rugose, but is thrown into ridges
forming an entostyle and a small protostyle. There is a cingulum and the tooth
bulges below it. Here, as in all other M2’, the posterior pillar projects less in a
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 433
lingual direction than does the anterior (cf. Hopefield 4024, Old. 3655). In
all other upper molars the posterior pillar projects more lingually.
There is a typical marked angulation of the lingual surface from the base
towards the apex in a buccal direction.
Occlusal surface. The central pits are most irregular on the lingual side,
the buccal enamel lip being V-shaped, while the lingual enamel is irregularly
U-shaped. The pits of the two pillars are in continuity in the region of the
mesostyle.
Roots. ‘Typical formation of the lingual root with its more massive anterior
portion formed by a vertical depression between the anterior and the posterior
portions of the lingual aspect. The buccal roots are roughly triangular in
shape, the posterior one being larger, and its tip curves posteriorly.
109 Olduvai BK II (1955) (plate 14(d), (e), (f))
Isolated upper left M? in early wear, with a portion of the hypocone
broken off. The enamel is finely rugose. There is a cingulum present and a
very small entostyle, while a ridge of enamel represents both the hypostyle and
the protostyle. There is a slight bulge below the cingulum and the lingual
surface slopes at a marked angle towards the apex in a buccal direction.
Buccal surface. 'The cingulum and its styles are typical of the teeth described
previously, except that the styles are not as obvious as, for example, in F 3655.
Occlusal surface. The central pits are in continuity with each other. The
paracone is rather widely separated from the protocone, but the metacone is
closer to the hypocone, and in fact the apex of the metacone tends to be twisted
in a lingual direction.
Roots. ‘The roots are broken off at the base.
F 2992 Olduvai II, in situ with ‘A’
This is an isolated upper molar in a very fragmented state. It is not
possible to determine which molar it is and to which side it belongs. It shows
characteristics similar to analagous portions of upper molars previously
described. It is in a fairly advanced stage of wear.
97 Old. BK II, 1952
Isolated right canine with the mesial portion of its enamel broken away.
The tooth is in an advanced stage of wear.
Buccal surface. Marked cingulum with a rounded bulge above it. The
enamel is coarsely rugose. The tooth is partially bilobed, the outer lobe pro-
jecting beyond the true transverse plane of the longest axis of the root.
Lingual surface. ‘The dentine is hollowed out laterally. The occlusal edge
shows a broad rim of wear. The occlusal edge of the tooth slopes outwards and
backwards.
Crown height ¢. 32 mm. Maximum breadth 15°O mm.
Length 28+ mm. Tooth height c. 60 mm.
434. ANNALS OF THE SOUTH AFRICAN MUSEUM
F 2991 Old. IS, 1941 (plates 18(a); 19(a); 20(a))
Isolated right P,, in a fairly advanced stage of wear.
Buccal surface. Coarsely rugose; marked cingulum especially at the base
of the hypoconid. A nodular ectostylid is visible.
Lingual surface. There is a cingulum. Rugosity is coarse, although the
tooth is worn smooth over the major part. The parastylid and the metastylid
are prominent, the parastylid rising up as a distinct ridge from the cingulum.
The median ridge of the metaconid is present.
Occlusal surface. The central pit of the anterior pillar is rather wide
anteriorly; it narrows in the centre, and then forms an open slit between the
metastylid and the entoconid. The posterior pillar has a central pit which has
a rounded formation anteriorly and which narrows posteriorly in the region
of the entostylid.
Roots. ‘They are broken off near the base, the posterior root appearing
large and quadrangular-shaped, the smaller anterior root being somewhat oval
with a concavity on its posterior aspect. On viewing the posterior aspect, the
‘apron’ effect of the enamel is visible.
2 Old. BK II (plates 15(a); 16; 17)
Isolated right P,. It has the general characteristics of F 2991, although
it is in a slightly less advanced stage of wear. The roots have been repaired and
replaced at an abnormal angle.
5 Old. BK II (plates 15(d); 16; 17)
Isolated left P,, with the roots broken away. The tooth has just com-
menced wear.
Buccal surface. It shows a coarse rugosity and a cingulum which is particu-
larly accentuated at the base of the hypoconid. An ectostylid is present on the
posterior part of the protoconid.
Lingual surface. Cingulum present. The entoconid shows a marked bulge
about half-way up the posterior aspect of the tooth. The median ridge of the
metaconid is prominent and there is a distinct metastylid which overlaps the
lingual surface of the entoconid and is separated from it by a wide interval
near the occlusal surface, but is fused with it near the base of the crown.
Occlusal surface. ‘Two wide central pits are present. The hypoconid is
separated from the protoconid by a wide V-shaped interval in the upper 3 of
the tooth, but is fused with it in the lower 4 (as in F 2991 and 4).
7 Old. BK I
Isolated right P,. The roots are absent. Part of the base of the anterior
pillar is missing, and part of the lingual surface is absent.
Buccal surface. Coarsely rugose, and there is a thick cingulum at the base
of the posterior pillar. Very similar to 5, except that the hypoconid is not
separated from the protoconid by a wide interval, but only by a furrow, and
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 435
that the ectostylid appears here only as a minute nodule. The tooth has just
erupted and is not in wear at all.
96 Old. BK II, 1952
Isolated left P,, with a part of the buccal surface, the base and the roots
missing.
Lingual surface. Very similar to that of F 2991.
Occlusal surface. Is also similar to F 2991 except that the central pit of
the anterior pillar is now closed off and there is a small island of enamel near
the parastylid, and the enamel of the entostylid has fused with that of the
metastylid.
105 Old. BK II ex. 1953 (plates 18(c); 19(c); 20(c))
Isolated right M,. Part of the anterior root is broken off and a portion of
the buccal enamel of the posterior pillar is chipped off.
Buccal surface. A small cingulum is present. The enamel is thickened in
the region of the ectostylid. The hypoconulid is formed by a marked vertical
ridge, although it is worn away at its base. The enamel is rather rugose.
Lingual surface. Small cingulum. It leads up to a parastylid on the anterior
pillar, and on the posterior pillar the entostylid seems to have a rather
broad base, and it is separated from the median ridge of the entoconid by a
slight vertical furrow. The base of the mesostylid can be identified and it is
continuous with the metaconid.
Occlusal surface. ‘The pillars have a rather circular shape. The central pits
are slightly curved with their convexities facing buccalwards, the anterior
portion of the anterior pit being larger than the posterior portion, while the
posterior part of the posterior pit is triangular and its lips are more widely
separated than those of the anterior portion of the pit. The buccal enamel of
the central pit of the anterior pillar is continuous with the lingual enamel on
the entoconid. There is a slight kinking of the enamel in the region of the
metastylid.
Roots. ‘The posterior root is broad and curves posteriorly at its tip in an
abnormal fashion and on its posterior surface it has a prominent ridge which
appears to be continuous with the unusually large hypoconulid. On the
posterior aspect of the base of the anterior root there is an aberrant root nodule
which has been broken off.
132 Old. BK II, 1953
Isolated right M,, in a very advanced stage of wear. Both roots are
broken off at the base.
Buccal surface. Rather rugose. Very slight cingulum. The crown is
rounded just above the cingulum. A small ectostylid is present. The anterior
surface of the tooth has a piece missing.
436 ANNALS OF THE SOUTH AFRICAN MUSEUM
Lingual surface. ‘The posterior pillar has almost all its enamel worn down
by abnormal wear (cf. specimen 93). The cingulum is barely recognizable on
the anterior pillar. ‘The enamel is rather rugose. The base of the central ridge
of the metaconid is just recognizable where it fuses with the metastylid.
Occlusal surface. ‘The central pits are irregular in shape. ‘The central pit
of the anterior pillar presents a bow-tie effect. ‘The posterior surface has its
enamel worn away completely.
95 Old. BK II, 1952 (plates 18(b), 19(d); 20(d))
A right M,. The roots are broken off at the base.
Buccal surface. There is a fairly well-defined cingulum on the anterior
pillar which is less marked on the posterior pillar, and a prominent protostylid
leads up from the cingulum. There is a small nodular ectostylid prolonged on
to the posterior pillar while there is an elevated ridge (hypostylid) on the
posterior aspect of the pillar. There is a small hypoconulid present. The
enamel is rather rugose.
Lingual surface. ‘There is a smal] cingulum. The parastylid is broken off
and the median ridge of the metaconid is prominent and rounded and is hardly
separated from the metastylid, so that the general impression of the anterior
pillar is that there is a very broad convex portion on the lingual surface (cf.
Old. 120, infra). On the posterior pillar, the entostylid is a small narrow ridge
and hardly separable from the entoconid. Seen from the anterior or the
posterior aspect, the buccal enamel presents the ‘apron’ effect.
Occlusal surface. The central pit of the anterior pillar is broad anteriorly,
while posteriorly it tends to slope towards the metastylid and in the central
portion the two enamel surfaces almost approximate each other. The central
pit of the posterior pillar presents a U-shaped appearance of the buccal lip of
enamel, and a V-shaped one of the lingual lip.
120 Old. BK II, 1955 (plates 18(e); 19(e); 20(e))
Isolated left M,, with most of the roots missing. It has a rolled appearance
and is in an advanced stage of wear.
Buccal surface. The pillars have a marked cingulum; on the anterior
portion of the posterior pillar, the cingulum is less marked, while it is most
marked on the talonid. A short ectostylid is present. The surface is rather
rugose. The buccal surface slopes quite markedly towards the apex in a
lingual direction. There is no or very slight rounding above the cingulum. A
slight cingulum is present on the anterior surface of the anterior pillar.
Lingual surface. Cingulum is marked on the pillars, but least marked on
the talonid. Leading up from the cingulum of the anterior pillar, there is a
rather marked parastylid; the metastylid is absent, and there is only a minute
entostylid. However, the central ridge of the metaconid is extremely broad
and rounded while the central ridge of the entoconid is also very large, but it is
smaller than the metaconid. Between the entoconid and the metaconid the
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 437
enamel folds rather deeply in a buccal direction emphasizing the metaconid
and the entoconid even more. The metastylid has fused with the metaconid
and thus produces this large bulge, and similarly part of the entostylid has
probably joined the entoconid to produce the latter’s large size (cf. specimen
95). The talonid is rather large and rounded; a hypoconulid is present. The
buccal enamel of the posterior pillar is continuous with that of the anterior
pillar but the two pillars are rather separated, which is a general variable
feature of the lower molars.
Occlusal surface. ‘The central pit of the anterior pillar is shaped like a bow-
tie, in that the anterior portion is triangular in shape, and the central pit is still
obvious, as also the posterior portion, but in between the two enamel lips are
lying against each other. In the posterior pillar, the central pit has a similar
appearance except that the anterior portion is more oval, the posterior portion
is narrower and is continuous with the central pit of the talonid, which is
fairly large. The enamel is generally very thick in this tooth.
Marsabit Road (plate 14(g), (4), (z))
This specimen has no number. It is a left third lower molar. Part of the
talonid is broken off, as well as a part of the anterior surface.
Buccal surface. The pillars have a marked cingulum and it is coarsely
rugose.
Lingual surface. Idem. The tooth is in advanced wear. The ridge of the
parastylid can just be made out, while the metastylid has fused with the
metaconid.
Occlusal surface. ‘The anterior central pit is L-shaped. The posterior
central pit is broadly U-shaped, and just behind it there is a rather wide
central pit on the talonid. Although the roots are broken, they appear to be
very short.
166 Old. BK II, 1955 (plates 18(d); 19(b); 20(b))
Isolated right lower molar, probably M,, in early wear.
Buccal surface. It has a distinct rounded cingulum, more prominent on
the anterior pillar than on the posterior one. The enamel shows a coarse
rugosity. The ectostylid is represented by a small ridge of enamel. There is a
small parastylid and a protostylid present. No hypoconulid. The metastylid
is very prominent.
Lingual surface. A well demarcated cingulum leads up to a marked para-
stylid and a slightly less marked entostylid which however is broken off near
the occlusal surface. The central ridge of the metaconid is well defined, and is
separated from the metastylid by a slight depression. The entoconid is broken
off.
Occlusal surface. ‘The central pit of the anterior pillar is slightly curved
and is wider anteriorly than posteriorly. The central pit of the posterior pillar
appears to be V-shaped: a part of the pit is broken away.
438 ANNALS OF THE SOUTH AFRICAN MUSEUM
116 Old. BK II Ex. 1953 (plates 15(c); 16(c); 17(c)).
Isolated right lower molar, just commencing wear, with the hypoconid
missing ‘and a portion of the entoconid broken. The roots are broken. It is
probably a M,.
Buccal surface. ‘The cingulum cannot be observed because the tooth has
been broken away. Surface coarsely rugose.
Lingual surface. The parastylid is a marked ridge. The central ridge of the
metaconid has a ribbed appearance, and a prominent metastylid is on the
upper half of the tooth; the central ridge of the entoconid also has a ribbed
appearance.
8 Old.
An isolated pillar of a left lower molar in early wear.
Buccal surface. There is a rounded cingulum present. Small ectostylid.
Rather rugose.
Lingual surface. Small cingulum leading up to a small ridged parastylid.
The central ridge of the metaconid is only obvious near the occlusal surface.
Occlusal surface. ‘The central pit shows an anterior widening and a posterior
narrow part, and the central portions of the enamel surfaces are continuous.
too Old. I Ex. 1953
This is an isolated entoconid of a lower right molar. Wear is just com-
mencing. It presents the general features of entoconids previously described.
98 Old. BK II, 1952
Broken isolated fragment of the entoconid of a lower right molar.
99 Old. BK II, 1952
An isolated fragment of a lingual pillar of an upper molar.
Marsabit Road
This specimen has no number. It is the posterior pillar of a right lower
M, with a smal]] fragment of its root. It is in advanced wear and has the
general characteristics of M, specimens previously described, except that its
buccal pillar is markedly twisted posteriorly. The central pit is V-shaped and
it has an irregular infolding of enamel posteriorly. There is a marked ecto-
stylid present.
The only measurements that can be taken are:
Maximum breadth cc. 32 mm.
Occlusal length 24:0 mm.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 439
Oldoway Surface (plates 18(/); 19(f); 20(/))
It is a right lower molar, probably M, in an intermediate stage of wear.
Most of the posterior pillar and root are absent, and the anterior portion of
the anterior pillar is broken away.
Buccal surface. Enamel rugose; cingulum is present and there is a slight
bulge above it.
Lingual surface. Fairly rugose, but worn smooth. The parastylid and the
metastylid are well defined.
Occlusal surface. Central pit has a typical bow-tie appearance.
Note.— The measurements of all the teeth are given in table 40 at the end of
section 2.
D. PosTcRANIAL SKELETAL REMAINS
116p Old. BK II, 1952 (plate 23(d))
The letter ‘p’ has been added to this number by the authors so as to
differentiate it from the dental fragment with the same number (supra).
Proximal end of a right ulna. This presents a massive olecranon process
which is separated from the articular facets by a massive rectangular ‘slab’ of
bone. The shaft of the ulna is broken off.
Olecranon process (posterior extremity) to articular facet
along superior border .. ee Ee we r .. 184 mm.
Olecranon process, maximum height .. Ae ae Lea Meritise
Olecranon process, maximum side-to-side breadth .. fo) 8 7G ER.
Breadth at centre of the ‘slab’ .. a se ee ae eG) | ROT:
Maximum breadth of articular process. . - i sO mm.
Most of its articular surface (i.e. about 4) is for articulation with the humerus;
only two small facets below this are for articulation with the radius. The
surface area of the radial articulation is relatively less than in the modern
giraffe.
115 Old. BR II, 1952 (plate 21(b))
Distal epiphysis of a right radius presenting a marked inferiorly projecting
tuberosity.
Maximum breadth at radial tuberosity See ee ae, ,b22 smi.
Maximum breadth at proximal end of fragment ~: 2) 195 mim.
Maximum A-P length .. “i ae = . Jc), 92: Tare.
341 Old. BK II, 1952 (plate 21(e))
Os magnum of the left carpus: very similar to that of modern giraffe.
Maximum length A-P .. ats ip oe ee relay igus sisi
Maximum breadth 4 < mat ae ae ie a Ge: onan:
Maximum thickness of postero-lateral side... ue Si f- SOy aaa:
440 ANNALS OF THE SOUTH AFRICAN MUSEUM
114 Old. BK Il, 1952 (plate 22(a))
Distal end of a metacarpal and a piece of the distal shaft: fragments have
been broken off, and the distal end has been chipped and rolled. In com-
parison with the distal end of the metatarsal (vide infra, specimen 314) the
shaft presents a definite flattened appearance, convex anteriorly and scooped
out posteriorly. It is almost identical in appearance to a specimen described
by Dietrich (1937), E 122 from Oldoway (his text-figs. 1 and 2, and table VI,
fig. 1), and another from Serengeti (Garussi-Korongo 1.39) also described by
Dietrich (1942) (his table XXII, fig. 187). The latter specimen is still a young
individual with an epiphysis; specimen 114 from Olduvai has a fused epiphysis
and. the distal end appears to be broader.
Maximum breadth across condyles as .. LOh yuan
Maximum breadth of lateral condyle, taken anheniorly .. ) RP aataa
Maximum breadth of medial condyle, taken aa «| 2 pana
A-—P length of the condyles ue a .. 54+ mm.
60 mm. above the distal end: A—P of the ee oe .. | 42 diag
Breadth of the shaft .. MNRAS (0920958011
100 Old. BK II, 1952 (plate 23(c))
Distal extremity of a right femur, consisting of 2 condyles and the patellar
condyle.
Maximum breadth across the condyles. . ae 3 .. SOT mags
Maximum breadth across medial condyle aie ie «| Ge staal
Maximum breadth across lateral condyle ve Me .. , 55), mama.
Maximum breadth across patellar condyle... ae +s 7G. aaa
Cord length of the patellar condyle in the centre... .. ‘TOG y mama
tor Old. BK II, 1952 (plate 21(c))
Distal end of a left tibia which articulates with numbers 102, 103. Adult.
It presents a marked bulge above the medial malleolus and another large
rough tuberosity on the antero-lateral aspect just above the articular surface.
Leading down from the shaft to this tuberosity there is a large linear ridge.
Distal extremity: Maximum A-—P bye Me up .»») (OR tomaame
Maximum breadth .. By Ne .» 20. ial
About 80 mm. from distal end: A-P_ .. ps “ . |) ear ca
Breadth Ai Wi, A) Gap aaa
112 Old. BK II, 1952
Distal extremity of a right tibia, which is a in many respects to IOI,
except that it is smaller.
Distal extremity: Maximum A-—P di a ii li CG, Mae
Maximum breadth .. oy ni a) TOG) maa
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 441
103 Old. BK II, 1952 (plates 23(a), (b); 24(c))
Left calcaneum, articulating with 102 and 104. It is a massive bone with
a markedly prismatic proximal tuberosity (tuber calcis). The body of the bone
is broad from front to back, and constricted from side to side. The facet
articulating with the posterior surface of the astragalus (talus) is irregular in
shape, and has a narrow downward and posterior projection, while it is broad
and quadrangular above this. The fibular facet, for articulation with the
fibular sesamoid, is quadrangular in shape, convex from front to back and is
angulated from behind forwards in a medial direction. At the distal extremity,
on the inferior aspect of the lateral side, there is a concave articular facet
arched upwards for articulation with the cuboid; it has a lateral convex
border, and medially it has a concave border.
108 Old. BK II, 1952
A left calcaneum, shorter than 103, presenting roughly the same features,
except (1) that the tuberosity is more rounded and massive, (2) that the body
is shorter, and (3) that the fibular facet is smaJler and more angulated medially.
103 108
mm. mm
Maximum length of calcaneum . oy 216 198
Maximum breadth (side-to-side) of tuberosity . . we 69 67
Maximum height (A—P) of tuberosity. . te 69 €. 62
Maximum length (A—P) opposite the fibular facet .. 89 gI
Body length from the superior border of astragalus facet
(along the anterior border) .. AP ai ‘oe 131 CEO
Minimum body breadth . F fis 39 42
Fibular facet: A—P length (on the convex portion) . 39 33
Breadth .. ae ue ar ay 28 24
TABLE 21
102 Old. BK IT, 1952
Astragalus (talus), belonging to the left side, articulating with the distal
end of the tibia No. 101 proximally, and with No. 104 distally, and with No.
103 posteriorly. The proximo-lateral articular ridge, for articulation with the
lateral fossa of the tibia, is large and wide, whereas the medial articular ridge
is narrow and has a large articular surface on its medial aspect for the medial
malleolus of the tibia. On the lateral aspect of the bone, just behind the mid-
point, there is a big oblique quadrangular-shaped surface for the articulation
of the calcaneum, and at the anterior end there is a small irregularly rounded
facet for articulation with the anterior extremity of the calcaneum. Between
these two surfaces there is a rough, hollowed-out region for the interosseous
ligament. On the anterior aspect, the fossa for reception of the lower border
of the tibia is long and saddle-shaped.
107 Old. BK II, 1952 (plate 21(a))
Right astragalus, with features similar to those previously described in
specimen No. 102. Because of the proximity of their discovery, they probably
belong to the same individual. Articulates with No. 110.
442 ANNALS OF THE SOUTH AFRICAN MUSEUM
102 107
mm. mm
Maximum proximo-distal length .. Bs 113 112
Maximum A—P diameter, medially Me 73 va
Maximum A—P diameter, laterally Be 64 63
Maximum breadth, proximally .. 4 87 86
Maximum breadth, distally a ae ¢. 75 76
Maximum articular breadth proximally .. 74 79
Maximum articular breadth distally ay ¢. 75 76
TABLE 22
104 Old. BK II, 1952 (plates 21(d); 24(g))
A left cubonaviculare, articulating proximally with astragalus No. 102
and calcaneum No. 103, distally with cuneiform No.:105 and metatarsal
No. 106. Proximally, the medial facet for articulation with the astragalus is
much broader than the lateral one, the two being typically separated by a
ridge. The tuberosity of the naviculare is short but very broad and massive.
Laterally, on the proximal surface, is the bean-shaped facet for articulation
with the calcaneum. On the distal surface, the articular facet of the cuboid
which articulates with the upper surface of the lateral metapodial is longer
and broader than the medial articular facet of the naviculare for the fused
cuneiforms. Posteriorly to this facet and continuous with it, there is a small
rounded facet for the external cuneiform. Posterior to the cuboid articular
facet for the metatarsal, there.is a well-defined groove which runs transversely.
Posterior to this groove there is a tuberosity which does not have an articular
facet for articulation with the metatarsal. This facet is present in the modern
giraffe.
109 Old. BK II, 1952
Isolated right cubonaviculare. The features are very similar to those
described in the previous specimen (104).
110 Old. BK II, 1952
Right cubonaviculare, articulating with No. 107, 110 A and 111. It has
identical features to No. 104. Probably belongs to the same animal.
104 110 args
mm. mm. mm.
Maximum breadth (side-to-side) across the centre 110. 108.) Ton
Maximum A-—P length across the tuberosity of naviculare 106 ~=6106 95
Maximum length of naviculare articulating facet for cuneiform 55 55 48
Maximum breadth of naviculare articulating facet for cuneiform a7 oF 35
Maximum length of cuboid articulating facet for metatarsal 61 61 55
Maximum breadth of cuboid articulating facet for metatarsal 45 48 42
TABLE 23
105 Old. BK II, 1952 (plate 21(g))
Left cuneiform consisting of the fused I and II cuneiforms. It has an oval
shape. It articulates with No. 104 and 106. The proximal surface is concave
from front to back. The distal surface is slightly convex in the central portion.
Medially and posteriorly there is an irregular prominence, the upper border
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 443
of which is in the same plane as the articular facet. On the surface of this
prominence are a number of almost parallel grooves which run from above in
a downward. and forward direction: they are probably grooves formed by the
astragalo-metatarsal ligament. On the antero-medial part of the superior
border, the articular surface has a small lip which projects downwards. It
articulates with a similarly shaped small projection on the cubonaviculare, at
the junction of the cuboid with naviculare, at the antero-medial part of the
opposing facet.
r10 A Old. BK II, 1952
A right fused cuneiform, articulating with No. 110 and 111. It is identical
in appearance to No. 105, but its posterior portion is broken off.
105 110A
Maximum A-P length Me oe ig sq 4 000mm. 59 mm.
Maximum breadth side-to-side... ae / o> 4am: 40 mm.
Maximum thickness (postero-lateral) a! se) Os Srna 200 mana.
106 Old. BK II, 1952 (plate 24(a), (b), (f))
This is a proximal end of a left metatarsal and a piece of the shaft. It
articulates with Nos. 104, 105. The anterior median groove is extremely wide
with marked ridges on each side. Proximally are three articular facets—two
are kidney-shaped facets for the cubonaviculare, while in between them and
slightly medially and posteriorly is the facet for the small rounded external
cuneiform.
111 Old. BK II, 1952
Proximal end and portion of the shaft of a right metatarsal, showing
similar features to 106. It articulates with Nos. 110, 110 A.
106 yigey |
mm. mm.
Maximum length (A—P) at proximal end 81 84
Maximum breadth at proximal end .. i pas Hi 93 95
About 60 mm. below: A-—P of shaft * a at 67 65
Breadth is a a oe 65 GP
Maximum A-—P of medial articular surface .. i “fi 59 59
Maximum A-P of lateral articular surface .. Mi aye 64. 58
Maximum breadth across centre of medial articular surface 33 36
Maximum breadth across centre of lateral articular surface 40 c. 50
TABLE 24
314 Old. BK II, Ex. 1953 (plate 24(d), (e))
This is the distal third of a right metatarsal. It presents a deep and wide
central groove anteriorly, of which the lateral lip is more prominent and
higher than the medial. The anterior surface has a general convex appearance,
while the posterior surface is flattened. Posteriorly a shallow and ill-defined
groove can be seen centrally, leading down to the space between the trochleae
(‘Rollen’, Dietrich, 1942). On the outer aspect of each trochlea, there is a
444 ANNALS OF THE SOUTH AFRICAN MUSEUM
fossa for the attachment of collateral ligaments, and the lateral one is deeper
and larger than the medial. Above the medial fossa there is a rough tuberosity
which is larger than the Jateral. The trochleae are big and separated by a
deep groove; at the base of the groove, where fusion has occurred, there is an
extensive central pit extending upwards. The grooves for the sesamoid bones
are shallow, the deepest one being the most lateral.
Maximum breadth at the trochleae .. bie He . 2) OG
Breadth of the lateral trochlea .. ae it iy el aie
Breadth of the medial trochlea .. bs oh se ty 5
A-P length across the trochlea, medially Ps oe 2) OT
laterally zs MAMem onan bu. (Oe,
Breadth of the distal extremity across the tuberosity .. ie Oe
Shaft some 120 mm. above the distal extremity
maximum breadth os ie Be iy 2 a
maximum A—P as ty He My, Me - 1. 6.00
M 14687
mm.
mm.
mm.
mm.
mm.
mm.
mm.
my.
This is an articulated hind limb in the British Museum (Nat. Hist.) and
Hopwood’s paratype (1934). ‘The femur, proximal end of the tibia and the
and and 3rd phalanges are missing. The measurements (mm.) are:
Tibia
Total length . sik ba bel . | AOOre
Length: crest af tibia — distal estiveanite dl, ul 2 r23e
Distal extremity: Maximum A-—P ae a 2 a
Maximum breadth yi oe +. 4 BOS
Metatarsal
Maximum length .. aN a se Lo) AE
Proximal extremity: Maximum A-P oe cp ape oy
Maximum breadth si. a MOS
Distal extremity: Maximum A-P .. Me a stay ge
Maximum breadth ae ie ahh AGO
Phalanx I
Maximum length .. si Be 1 Deg '1 0)
Proximal extremity: Maximum A-P srk a py sane 5,0)
Maximum breadth .. mn 5
Distal extremity: Maximum A-P .. a hee Ne pec},
Maximum breadth ayy fe anne
113 Old. BK II, 1952
185 Old. BK II, Ex, 1953 (plate 22(c))
F 364 Old. S I, 1941 (plate 22(b))
Old. ‘Surface’ (plate 22(d))
F 3297 Old. II, 1941, with ‘A’
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA A445
F 365
These are all proximal phalanges and it is not possible to say with any
degree of certainty whether a particular phalanx is a medial phalanx of the
right limb or whether it belongs to the lateral side of the left limb; or whether
a lateral phalanx of the right may belong to the medial side of the left. Further-
more, it is not possible to state whether an individual phalanx belongs to the
fore or hind limb; but we have observed that in one particular extant animal
the proximal phalanx of the fore limb is more massive than that of the hind.
limb. On this basis it is suggested that specimen F 3297 and ‘Oldoway surface’
probably belong to fore limbs, while the other four specimens probably belong
to hind limbs. ay
185 113 HOLA StinfacthimiaZ297. F365:
Maximum length .. Ho P07 108 113 120 — 114
Breadth (side-to-side) at base as 50 ¢. 51 44 60 60 52
A-P at base. ae We 54 C. 51 48 - 58 57 51
Minimum breadth, ‘shaft .. ks 46 43 Ba 47 — 39
Distal extremity:
Maximum breadth (side-to-side) 48 49 46 58 — 48
Maximum A-P length .. oe 33 34 29 38 — 31
TABLE 25
342 Old. BK II, 1952 (plate 21(/f))
Sesamoid bone articulating with the head of the middle phalanx and the
base of the distal phalanx.
Maximum A-—P 53 mm.
Maximum breadth 36 mm.
Maximum thickness 34 mm.
CHAPTER 2
ORANGE FREE STATE (UNION OF SOUTH AFRICA)
A. LOCALITIES
Fossil Sivatheriinae have been recovered from five different localities of
the Orange Free State which extend over a large area (the furthermost points
being about 150 miles apart), but they all belong to the Vaal River basin
(fig. 18). Consequently, in spite of the fact that some of the specimens have
been described by different individuals, that they have been found at various
stratigraphical levels and that they are housed in different museums, the
general geological picture of the Vaal River basin (Cooke, 194.9) provides good
reason to consider them in one group.
The fossil specimens are recorded as being derived from the following
Sites:
1. MMK 3685 (McGregor Memorial Museum, Kimberley) is stated as
aw
446 ANNALS OF THE SOUTH AFRICAN MUSEUM
gs PRETORIA
£7) m{LICHTENBURG
we Pi
JOHANNESBURG
VEREENIGING
Cornelia
@ KROONSTAD
@ Hoopstad
Viakkraal
and A Florisbaa \?
Douglas ;
Soe :
a
~ 5
Se
=o
to]
BLOEMFONTEIN
ai- Waldeck’s Plant
ac Gong-gong
a3- Barkly-West
Fic. 18. Map of Vaal River basin indicating major fossil sites (A)
(modified after Cooke, 1949).
coming from an ‘unknown locality of the Vaal River basin’ (Haughton, 1922).
Assigning this tooth to a new genus and species, Griquatherium cingulatum,
Haughton stated that it came from the collection of Mr. A. Grumpelt at
Barkly West, and Cooke (1949) stated that on the basis of the other specimens
in this collection, it is quite likely that the Griquatherium specimen came from
the 60-foot terrace at Waldeck’s Plant or Gong-gong.
2. In 1926, three Sivatheriinae teeth were recovered from the upper
layers at Florisbad (Dreyer and Lyle, 1931). The actual teeth were sent in
1932 to scientists in Europe for study, but unfortunately they were not described
and it is now not possible to trace them. The only remnant of the specimens
is a poor plaster-cast of the crown of a lower molar (C 1492) which is housed
in the Nasionale Museum, Bloemfontein.
Notes on the geology of Florisbad have been published on various occasions
(Dreyer and Lyle, 1931; Dreyer, 1938; Hoffman, 1953; Oakley, 1954a; Singer,
1956, and Meiring, 1956). Oakley (1954a, page 84) states:
‘The Florisbad deposits consist of sands, intermittently ejected by springs
of gaseous water during Pleistocene and recent times, alternating with seams
of peat formed by salt marsh vegetation which spread across the area when the
springs were quiescent. There are two parallel lines of spring centres (or
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 447
eyes’) which have become sealed off progressively in an easterly direction.
Thus the “‘eyes’’ are fossilized on the western side of the site but still active on
the eastern side.
‘Fossil mammalian bones and teeth and Stone Age implements occur in
the beds of sand formed by the spring waters during Pleistocene times. Owing
to the occasionally disruptive action of the springs, when old “eyes”? are
reopened, it is not always possible to be sure of the original stratigraphical
position of specimens found in these deposits. The most reliable finds are
those from below uninterrupted seams of peat.’
Despite deficient data (Dreyer and Lyle, 1931, p. 5, and the Nasionale
Museum Register), Oakley (1954a) seems to have obtained information some-
where that a tooth of the extinct Sivatherine (Orangiathertum) was found
between Peat II and Peat III.
3. A horn-core fragment, the type specimen of Orangiatherium vanrhyni v.
Hoepen, was merely mentioned by van Hoepen (1932). He also mentioned ‘a
terminal fragment of a large antler and a series of large teeth, which were
probably associated’. These specimens, presently housed in the Nasionale
Museum and registered under the numbers C 431A, C 431B, and C 426
respectively, are derived from the farm Tierfontein, on the Vet River, 9 miles
from Port Allan (personal communication from the Director of the Museum,
Dr. A. C. Hoffman) (fig. 18).
4. Specimen F 39 (Archaeological Survey of the Union of South Africa,
Johannesburg) is stated by Cooke (1949) to be derived from ‘an unknown
locality in the Vaal River basin’. This is the type specimen of Griguatherium
haughtont Cooke.
5. Two Sivatherine upper milk molars were identified in the collection
of specimens from Cornelia, in the Nasionale Museum, Bloemfontein, by one
of us (R.S.): these specimens are not registered and are allocated B! and B?
by the authors.
The Cornelia site consists of a large erosional area adjacent to a small
branch of the Vaal River. Specimens are found on the floors and on the sides
of “dongas’ (eroded clefts) which are washed by seasonal rains and partly
covered by flooding of the nearby river, and consequently there are numerous
redepositions. The surface consists of hardened calcited sand which rests on
varying projections of Karoo formations. A typical view of the site is shown in
plate 5 of Oakley’s above-mentioned paper (1954a).
B. List oF MATERIAL AND DESCRIPTION
MMK 3685 : Isolated left M*— Vaal River (? Waldeck’s Plant).
1492 : Cast of occlusal portion of left M,—Florisbad.
Cc 426 : 4 isolated upper molars (right M? and M®, left M, and M?)—Tierfontein.
C 431 : Left posterior horn-core and a right (?) anterior horn-core (ibid.).
F 39 : Stated to be an isolated anterior pillar of a lower left M, or M,;— Vaal
River (unknown locality).
B! and B? : Isolated right DM® and DM*— Cornelia.
448 ANNALS OF THE SOUTH AFRICAN MUSEUM
I. HORN-CORES
C 431
There are two specimens marked C 431 from Tierfontein. The one is an
almost complete posterior horn-core with the tip missing, and this is here
designated C 431 A. The other, C 431 B, is a fragment of an anterior horn,
and not a part of A.
C 431 A (plate 29(¢), (d))
This is a left posterior horn-core (vide infra, ‘Discussion’), the base of
which is pear-shaped, the narrower portion being anterior. The apex of
the hollowed-out portion of the base, formed by sinuses, is 27 cm. distant from
the broken edge of the base. From the base, the anterior border runs out- —
wards in a gentle arc towards the first knob, then upwards and slightly medially
and then laterally, giving the broad surface of the horn a double twist.
The horn is flattened from side to side opposite the first and second knobs,
but above these it tends to bulge where the anterior border becomes
rounder.
Grooves on antero-medial surface (cf. Old. 3.53)
From the base there are three fairly deep and broad grooves starting near
the front at almost a single point and running obliquely up and back at an
angle of about 45°. The most medial groove ends in a trifurcation at the
posterior border opposite the first knob. Along the antero-lateral border, at
the base of the broken-off flange, three deep grooves pass vertically up parallel
to the border. Just below the first knob they tend to deviate from each other,
the anterior one running along the anterior border and, passing behind (lateral
to) the second knob, it divides into a number of smaller grooves running up
almost parallel to each other towards the tip.
Between the levels of the first and second knobs there are four other deep
main grooves on the convex surface which run up this surface fairly parallel to
each other. The posterior one divides into three just below the level of the
second knob, and these then pass towards the posterior border and run
laterally.
The anterior border in the region of the flange has two irregular rough
tuberosities with numerous vascular foramina. There are, in all, three knobs,
very crinkly in appearance and fairly evenly spaced from each other. Between
them are two small raised irregular tuberosities.
Circumference at base a ie (a ett Ne! 360
Circumference between flange and knob 2 at e..8Alh
Circumference 100 mm. above knob2 .. 258
Circumference at knob 3 .. a Se ey 255
Circumference at tip ay alls ave
Total length hh Ne a af 5 570+
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 449
A-P Side-to-side
Base .. cp Le Ria galt Ge oF
Between flange and knob 2 es oe ee 137 79
Above knob 2 (100 mm.) .. ap uy a, 93 7p
At knob 3°... fA ae se eile ae 97 65
At tip a; ne i Re ai ae — —
TABLE 26. Measurements of C 431 A (mm.).
C 431 B (plate 20(a), (d))
This is an anterior horn, probably right (if the grooves are taken to be on
the medial side). The base is flat from side to side and rather triangular with
the broad end posterior. Hollowing-out the base are three cranial sinuses.
The medial surface is extremely irregular with numerous ridges formed by
deep grooves. The grooves are deeper anteriorly and commence at the base
of the anterior border and pass in an inverted triangular fashion, the anterior
grooves being vertical and the posterior oblique. The anterior groove branches
about half-way up. The posterior part of the medial surface is roughened by
small, shallow grooves. ‘The anterior border has a knob just above the base—
it has a cauliflower appearance and is distorted by (? vascular) grooves. ‘The
top of the anterior border has another similar knob. The superior surface is
very irregular and grooved, the central portion being smooth. The posterior
border is concave, the upper end passing backwards and there is one small
protuberance at the base and one at the upper end. The outer surface is
rather smooth with a single deep groove near and parallel to the anterior
border.
Circumference at base ae ae Hs 280
Circumference at tip ae oe: Be 320
Total length: Anterior .. ae tr 62220
Posterior .. Fe Vo neGn LO
A-P Stde-to-side
Base... Me a oe Be WNT c. 70
Middle (opposite knob) ee Tol 55
ALT OMNP oe Be ae 123 61°5
TABLE 27. Measurements of C 431 B (mm.).
2. TEETH
MMK 3685 (plate 25)
An isolated upper molar in a medium stage of wear, with the roots broken
off near the base. This is the type specimen of Griquatherium cingulatum Haugh-
ton. Although it was originally described as a second molar (Haughton, 1922),
and stated by Cooke (1949) to be either a M? or a M3, it is here considered to
be unquestionably a third molar because of the relative decrease of the lingual-
ward projection of the posterior lingual pillar compared to the anterior one
(vide infra).
Buccal surface. Each pillar has a marked cingulum, that of the anterior
pillar leading to the broad base of the rounded marked parastyle. The central
median costa of the paracone is fairly distinct, the bulge commencing about
half-way to the occlusal surface.
450 ANNALS OF THE SOUTH AFRICAN MUSEUM
The mesostyle is distinctly prominent, projecting out markedly in an
anterior direction from the surface of the pillar and having a broad base
continuous with the cingulum of the posterior pillar on the one side and with
the cingulum of the anterior pillar on the other side, though in the latter there
is a slight vertical groove partly separating them. The vertical median costa
of the metacone is narrow and hardly prominent, commencing near the base
just above the rolled edge of the cingulum. Posteriorly, the cingulum of the
posterior pillar is continuous with the bulging base of the short metastyle.
Lingual surface. ‘The enamel is rather rugose. On the anterior pillar there
is a marked rounded cingulum, the lower border (the one towards the occlusal
surface) of which increases in thickness on the anterior surface producing an
unusually long, rolled and ridged protostyle. The cingulum of the posterior
pillar is relatively small and aimost absent in the region of the entostyle. On
the posterior surface of the posterior pillar the actual cingulum remains small
and ridged but a distinct elongated crest forms an unusual hypostyle. The
lingual surface slopes towards the occlusal surface at an acute angle from the
cingulum, but near the occlusal surface the angulation changes and the lingual
surface tends to become slightly more vertical. The slope of the buccal surface
on the other hand is almost vertical.
Occlusal surface. The central pits have widely separated enamel surfaces,
the pit of the anterior pillar being acutely V-shaped and closed anteriorly by
the anterior limb of the protocone. The central pit of the posterior pillar has
a more obtuse V-shape, being closed posteriorly in the region of the metastyle,
but open anteriorly and continuous with the space between the enamel surface
of the contiguous side of the protocone and the hypocone. The occlusal
surface of the posterior pillar is abnormally longer than that of the anterior,
the ratio being 114.4 (table 28).
Stvathertinae
Measurements (mm.) and indices M? M? MME
(Mean) (Mean) 3685
Occlusal length posterior pillar ee eee Bao 1144
Occlusal length anterior pillar ry
Maximum breadth posterior pillar ele 99°1 gt git
Maximum breadth anterior pillar
Maximum tooth length if a ae 47:6 515 53°8
Maximum tooth breadth .. is re 49°7 48°7 54°6
TABLE 28
From the table, the ratio of the breadth of the posterior pillar relative to
that of the anterior pillar corresponds closer to that of four other Sivatherine
M? than to that of two Sivatherine M?. From observations on the extant
material this ratio in M$ is a very constant one, and consequently, despite the
fact that the ratio for the relative occlusal length appear nearer to that of M?,
the authors consider that MMK 3685 is a M3. The length and the breadth
of the tooth are at the outer limits of the range of Sivatherines (tables
26, 40).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 451
C 1492 (Cast)
This is a plaster reproduction of the occlusal portion of the crown of a
left M,. Observations on the cingulum are impossible and what can be made
out from the enamel of the cast, it appears to be fairly rugose. The tooth does
not appear to be in an advanced stage of wear.
Lingual surface. There is a prominent metastylid and the buccal enamel of
the metaconid is continuous with the lingual enamel of the entoconid. The
parastylid is slightly prominent. The central costa of the metaconid appears
rather flattened.
Occlusal surface. ‘The anterior pillar is more rounded than the posterior
pillar on the buccal aspect; the central pit of the anterior pillar is ill-defined,
but that of the posterior pillar shows a fairly wide central portion, narrow
anteriorly and broad posteriorly.
C 426
This number is given to four upper molars. The authors have sub-
divided them into A, B, C and D which will be entered in the register of the
Nasionale Museum, Bloemfontein. A and B belong to the same individual
because of the obvious contact surfaces.
C 426 A (plates 26(a); 27(a); 28(a))
This is a right M? in an extreme degree of wear. Part of the paracone is
missing.
Buccal surface. A cingulum can be seen, especially in the region of the base
of the metastyle where it is considerably heaped up. There is no cingulum in
the region of the base of the mesostyle and the surface between meso- and meta-
style has been hollowed out, in the same ‘W’ formation as other teeth described
previously: there is a slight bulge in the region which leads up to the apex of
the metacone. The paracone is almost completely absent.
Lingual surface. Finely rugose. On the hypocone the rugosity is extremely
fine with additional transverse striations. On the anterior aspect of the proto-
cone the enamel is raised slightly at one spot, but the rest of this surface is
extremely smooth due to contact pressure, and this wear has even extended on
to the base of the root in this region. On the hypocone there is a very slight
cingulum. On the protocone, the cingulum is slightly more marked, but still
negligible. On the protocone too, the lingual surface bulges slightly above the
crown-root junction, and it can be seen to slope towards the apex in a buccal
direction. The posterior surface of the hypocone is worn right down to the
crown-root junction, only a small piece of enamel being visible here. Here
also, the wear has extended on to the root.
Occlusal surface. Despite the marked wear, the central pits are obvious,
the enamel edges of each pit of the anterior and posterior pillars being separated
to a fair degree. In the anterior pillar, the pit has an L-shape, the upright of the
‘L’” extending right up to the parastyle, which is broken off. However a small
452 ANNALS OF THE SOUTH AFRICAN MUSEUM
island of enamel belonging to this pit can be seen extending towards the meso-
style, but it is separated from the L-shaped portion by dentine which is
hollowed out by wear. The pit of the posterior pillar is irregular in shape, the
lingual aspect having its enamel thrown into two folds which project into the
centre of the pit, and the extremities of the pit extend towards metastyle and
mesostyle.
The dentine is particularly hollowed out between the enamel of the lingual
surface and that of the central pit. On each occlusal surface can be seen
striations which in some places are almost distinct scratches indicating a side-
to-side chewing movement of the jaws. The shape of the enamel of the lingual
surface viewed from the occlusal side is arc-shaped for both cones, the arc of
the hypocone being more flattened than that of the protocone.
Roots. Despite the marked fragmentation, it is possible to identify three
roots. The lingual root being composed of two massive pillars joined by a
plate, and the anterior pillar being the larger. The posterior buccal] root is
triangular in shape, while the anterior is broken off at its base and fragmented
so that its shape cannot be identified.
C 426 B (plates 26(b); 27(b); 28(5))
This is a right M? which is very fragmented, so that only a portion of the
enamel of the hypocone on the lingual surface is present, while that of the
protocone is absent. Both the metacone and the paracone are missing.
Lingual surface. Rugosity is fairly marked in parts, and there is a small
cingulum which tends to have a rounded bulge below it. The anterior surface
of the protocone shows some ridging of the enamel just below the small cingu-
lum (protostyle) while the rest of this surface, which is only represented by a
small fragment, is very smooth due to contact wear. A small piece of the
central pit of the anterior pillar is present, the enamel of the two sides of the
pit being separated to a fair degree. The roots are represented only by a portion
of the lingual root.
C 426 C (plates 26(d); 28(c))
A left M? which is a badly fragmented tooth with only protocone and
hypocone and a portion of the lingual root present.
Lingual surface. Rather rugose, with a poorly defined cingulum on the
hypocone and a well-defined cingulum on the protocone. On the lingual
surface of the protocone, near the occlusal surface, is a ridge of enamel parallel
to the cingulum. There is no bulge above either cingulum. The lingua] surface
of the protocone slopes only slightly towards the apex, while that of the hypo-
cone seems to have a slightly more marked slope. The shape of the lingual
enamel of the occlusal surface is arc-shaped for both cones. On the anterior
surface of the protocone the enamel shows a slight horizontal ridge (proto-
style) but most of the surface is smooth by contact wear. The enamel of the
hypocone ends abruptly at the posterior surface and this surface is hollowed
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 453
out and appears to be worn smooth by contact with the adjacent tooth. This
appearance is not uncommon in giraffids.
Occlusal surface. ‘The hollowed-out pits are irregular in shape, the enamel
of the lingual and buccal lips of the pit being fairly separated. Because of the
absence of parastyle, mesostyle and metastyle, it cannot be determined whether
there are any separate islands of enamel in those regions which may have been
linked to the central pit at an earlier stage of wear.
Roots. Most of the lingual root is present and it has a similar appearance
to that previously described for 426 A and B.
C 426 D (plates 26(c); 28(d))
A left M! which is very incomplete, only a portion of the protocone
remaining below a very fragmented lingual root. There is a small cingulum
and a very slight thickening of the enamel just below the region of the cingulum
where the tooth tends to bulge. The general shape and appearance of the
protocone is identical to that of C 426 C.
F 39 (plate 27(c), (d), (¢))
This is the type specimen of Griquatherium haughtont Cooke 1949. It has
been described as an isolated anterior pillar of a lower left molar, either M, or
M, (vide infra). It is in an early stage of wear. Although the evidence for the
viewpoint that this is half a lower molar is reasonable, peculiarly enough the
specimen also presents a number of features which raise considerable doubt
of the accuracy of this diagnosis and which could support the proposition that
this specimen is an upper premolar. For this purpose it is necessary to compare
it directly with Hopefield 4025 and Olduvai F 2989 (plates 45, 15-17). This
matter is discussed in detail below, but the authors consider the
evidence to be in favour of a left upper premolar and will describe the specimen
on that basis as follows.
Buccal surface. ‘The tooth is broken probably just below the crown-root
junction, and a portion of the posterior aspect is broken away. The rugosity
is fairly coarse. There is a marked rounded parastyle which is separated from
the prominent median costa by a broad groove, and the costa in turn is
separated from the prominent metastyle by a narrower groove. Near the
occlusal surface, the metastyle swings posteriorly in an arc.
Lingual surface. It is coarsely rugose. ‘The tooth is fractured at the crown-
root junction and a small rolled cingulum can be seen at the base of the posterior
lingual aspect. This appears to be sufficiently localized to warrant being called
an entostyle. From the anterior aspect, it is obvious that the enamel of this
surface presents a marked ‘apron’ effect. The lingual surface slopes markedly
downwards in a buccal direction to a point approximately at the junction
between middle and lower 4 of the tooth. Then the surface tends to slope more
vertically to the occlusal surface, so that the general effect is that the upper 2
of the tooth has a marked lingual bulge. This lingual bulge is the most promi-
454 ANNALS OF THE SOUTH AFRICAN MUSEUM
nent observed in the whole series of African Sivatheriinae. This matter will be
more fully dealt with in the discussion, but it is here noted that an X-ray of
the tooth (plate 27(c)) indicates internal vertical fracture lines, which may
have been caused by intrusive compressing breccia. This may be an explanation
for some of the excessive bulging observed.
Occlusal surface. ‘The central pit is fairly wide and open posteriorly, while
anteriorly the two enamel surfaces are in continuity with each other.
Posterior surface. A large piece of the enamel near the buccal aspect is
broken away, especially from the upper 2 of the tooth. But sufficient of the
enamel is present near the occlusal surface to ‘close’ the lingual and buccal
enamel crescents of the protocone. ‘There is also sufficient enamel on the
lingual surface to indicate that it is extending without interruption well towards
the buccal aspect: this is further back than would be expected if the enamel
were to be continued on to the contiguous surface of another pillar (on the
alternative supposition that this may have been a lower molar). The enamel
is heaped up slightly in the region of the protostyle.
Morphologically it may however be compared favourably with the
anterior pillar of M, and M, of Olduvai 92 (BK II, 1952). On the other hand,
if the probability of the dimensions of F 39 is calculated by the ¢ test in respect
of its being a M, or Msg, the following is obtained:
On the basis Value of P
of P39— iM, F39—2,
Occlusal length of anterior pillar .. 05 “I
Maximum breadth of anterior pillar “4 =)
Occlusal breadth of anterior pillar. . zo) 6
Buccal height of anterior pillar... Ol Ol
Because of the large dimensions of this ‘pillar’ (on the alternative supposi-
tion that it is a lower molar), the available measurements have been utilized
to reconstruct the whole tooth by a comparison with M, or M, of other known
Sivatheriinae. Calculated on this basis, the Tr./A—P index of F 39 would
be:
M, M;
F 39 de ee ih 641 46-1
Mean of Sivatheriinae .. 70:0 49°7
. Occlusal length of anterior pillar
and the ratio ——————s—— —____—_—_—_ i VOOn:
Maximum breadth of tooth
F 39 oe site ake aE 812
Mean of 12 Sivatheriinae M, .. 74°5
Mean of 10 Sivatheriinae M, .. 752
Consequently, it is reasonable to state that one cannot assign F 39 as a definite
M, or M;: it is furthermore clear that from the point of view of the length
and breadth of the tooth (table 29), it is unlikely to be an anterior pillar of a
lower molar. Its height is greater than that of any tooth in the available series
(see also ‘Discussion’).
5
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 455
Anterior pillar: Maximum tooth Anterior pillar: Maximum tooth
Stvatheriinae occlusal length length maximum breadth breadth
M, (mean of 12 specimens) ne. 26-0 512 34°9 35°7
M, (mean of 10 specimens) Bi 25°0 68-6 33°3 33°6
F 39 Actual measurements Ae 32°5 c. 40
Inferred measurements:
M, si ue 64:0 41-0
If M, ma *, 87°5 40°3
TABLE 29
B! and B?
These two teeth are both from one site, probably Cornelia, but there are
no records available concerning their discovery. The one tooth B! has three
distinct roots, one lingual and two buccal, and is therefore an upper molar.
In B? the roots are broken away and the tooth is only just commencing wear
on proto- and paracone. It is also obvious that both teeth belong to the same
jaw and the same side because of the exact fit of their contiguous surfaces.
Because of, first, the small size of the root in B!, secondly, of the hollowness of
the root and the tooth, thirdly, of the thinness of the cement and enamel, and
fourthly, of the general appearance of B! and B?, it is considered that these
teeth are deciduous. Consequently, B! is diagnosed a right DM? and B? is a
right DM+.
B* (plate 30(2), (¢); (¢))
Right DM. It is in early wear
Buccal surface. Small cingulum present. Rugosity very fine. Metastyle,
mesostyle and parastyle have a very obvious rib-like effect, the parastyle being
the largest of the three styles and being decisively rounded and separated from
the lingual surface of the paracone by a distinct cleft. The ridge leading up to
the apex of the paracone is more obvious than that leading up to the apex of
the metacone, which is almost absent. The appearance is typical of the Family,
the surface having an undulating effect, especially when seen from the lingual
or the buccal aspect, the crests being formed by the apices of the paracone and
protocone, and of the hypocone and the metacone, while the depressions are
opposite parastyle, mesostyle and metastyle.
Lingual surface. ‘The protocone has an obvious cingulum, while that of the
hypocone is less obvious. On the antero-lingual aspect the enamel is heaped
up and forms a ridge (protostyle). Similarly on the postero-lingual aspect of
the hypocone, there is another ridge of enamel (hypostyle), but this is very
small and less extensive than the protostyle. The surface is finely rugose. Just
below each cingulum, there is a bulge which is less marked on the hypocone
than on the protocone, and from this bulge the lingual surface slopes fairly
acutely towards the apex. From the anterior and posterior aspects, the crown-
root junction has an irregular line which is ‘apron-like’, as in the Hopefield
upper teeth.
Occlusal surface. ‘The protocone is quite separate from the hypocone but
the central pits of the two pillars are continuous opposite the mesostyle, and it
456 ANNALS OF THE SOUTH AFRICAN MUSEUM
can be seen that the separation of the pits would have been caused at a later
stage of wear by the hypocone going to meet the paracone to form the meta-
conule. The lingual and buccal lips of the pits are widely separated, maximally
at the centre of each pillar. The meeting of the protocone and the paracone
takes place at the parastyle where their dentine and enamel surfaces are
continuous. But the meeting of the hypocone and metacone is only by contact
of their enamel surfaces.
Roots. The lingual root is rather small and oval-shaped, the separation
into the two thickened portions of the root being hardly noticeable. The
cement is very thin, the root and the tooth being completely hollow. The two
buccal roots are thin and hollowed although the cement is thicker than in
the lingual root; the posterior one is triangular and the anterior one is oval in
shape.
B® (plate 30(b), (4), (f))
Right DM?.
Buccal surface. ‘Vhere is a slight cingulum present on the hypocone and it
leads up to a very marked mesostyle and a slightly Jess marked metastyle. The
parastyle is prominent and rounded, and it has a small nodular excrescence
on its buccal aspect. ‘The mesostyle is somewhat split by a vertical furrow.
The buccal aspect of the paracone is much more marked than that of the
metacone ‘The buccal surface of the metacone is distinctly spatulate-shaped
as is commonly found in milk dentitions.
Lingual surface. Finely rugose. The hypocone has a slight bulging cingu-
lum from which the lingual surface slopes rather sharply to the apex. Although
the protocone is rather fragmented at its base, there seems to have been no
cingulum, and the lingual surface also slopes rather sharply towards the apex.
On the anterior surface, the enamel is thrown into a small ridge (protostyle)
just above the crown-root junction.
Occlusal surface. ‘The hypocone and metacone are just beginning to wear,
whereas the protocone and paracone are worn to a more marked degree. All
four cones are separated, their fusion only occurring between the hypocone
and metacone near the metastyle, and between the protocone and paracone
near the parastyle, while paracone and metacone are almost completely fused
on the buccal surface medial to the mesostyle. Consequently, the central pits
are ‘wide open’ although the two cones of each pillar have fused at the base
of each pit at the base of the tooth.
Roots. No roots present. They have been broken off. The tooth is
hollowed out.
Shape. The protocone is V-shaped and the hypocone is slightly more
arched and broader. The paracone is almost a straight line while the metacone
is spatulate-shaped.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 457
CHAPTER 3
MAKAPANSGAT (NORTHERN TRANSVAAL, UNION OF SOUTH
AFRICA)
The various discoveries in and the geology of the Limeworks Cave in the
Makapan Valley have been fully described by Dart (1954), Oakley (1954b),
Brain, van Riet Lowe and Dart (1955), Howell (1955), Brain (1957, 1958),
and Wells and Cooke (1957) in whose publications further references may be
obtained.
All the material described here is on loan from the Bernard Price Institute
for Palaeontological Research (Johannesburg), and is derived from the Lime-
works dumps wherein out of 1,862 skull remains already recovered some 30
belong to Giraffids (Dart, 1957). However one specimen, M 553 B}, is from
an unknown locality in the Makapan Valley, ‘and probably from one of the
limeworks’ (Cooke and Wells, 1947). Some of the material is still heavily
filled with calcified grey cave breccia.
The material received for study may be divided into two major groups:
the first group consists of giraffid teeth referable to Giraffa, the determination
of the species of which will be mentioned later. The second group is composed
of fragments of the dentition and of the jaws of Sivatheriinae. Both groups were
recovered from the same deposits, and from the nature of the breccia it is
probable that they were contemporaneous in so far as the limits of the formation
of the breccia are widely separated by a large period of time.
List oF MATERIAL
A. GIRAFFA
(1) Upper dentition
(a) Milk M 646 ae Pa) aright Py Nie
M 944 es. o2, «left DM?
M 536 we os leit: DM®
M 263 - .. fragment of right maxilla with DM?— M1?
M 533 a3 oe IOI
M 535 =: ~ fete DM?
MELE TS 9 5. a= left. DM?
(b) Adult M 532 se seit lett Re
M 531 De a! lett Re
M 264 ae “2 tebe PS
Merri. (o.. 20 gishit Pe
M 552 i gee 5 Eiht Ve
M551 at 6. dere Vit
M 550 Se .. fragment of maxilla with left M2?
M 528 Bie .. fragment of maxilla with left M?— M3
(2) Lower dentition
(a) Milk M 939 ey, .. fragment of left mandible, with DM, and DM,
M 540 a .. fragment of right mandible, with DM,— DM,
(6) Adult M 936 right M,
M 942 and Mit 13 joined fragments of right M,
M 938 3 .. fragment of right mandible with M,
458 ANNALS OF THE SOUTH AFRICAN MUSEUM
B. SIVATHERIINAE
(1) Upper dentition
Milk M 937 eh .. isolated left DM? or DM?*
M 524 bis .. right, probably DM?
M 941 i .. right, DM? or DM?
(2) Lower dentition
(a) Milk M 525 ae ie ete va.
M539B .. .. fragment of right mandible with DM,
(b) Adult M 527 of ond) LMICISOF,
Miri. se in) MCISOL
M553A .. .. fragment of left mandible with P,
M553B OL. .. fragment of right mandible with P,— P,
M553 BI .. .. fragment of left mandible with M,—M,
M 943 om isolated pillar of right M,
(See also Appendix. )
DESCRIPTION
A. GIRAFFA
The general appearance of these teeth is the same as that of the extant
Giraffa camelopardalis and nothing can be gained by describing each specimen
in detail. The only variations that may be noted are minor individual ones,
namely, those of irregularity of enamel ridges present in some and not in
others, the presence of entostyle and parastyle, of ectostylid and parastylid and
other variations which are also found in the non-fossilized extant material.
OM2 Makapansgat
es COPE C immature)
pM3
om4
DMo
DM3
DM,
5 10 15 20 25 30 35 mm
Fic. 19. Dimensions of deciduous teeth of Makapansgat Giraffa plotted against ranges
of variation in modern Giraffa camelopardalis.
—— = A-P length. ------ = Transverse (breadth). . = Makapansgat specimens.
(1) UPPER DENTITION
(a) Milk teeth
M 646 and M 944 (plate 31(a), (7), (9); 31 (6), (4), (7)
These are deciduous upper second molars, 944 being left and 646 being
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 459
right. They are in the same stage of early wear and are identical to each other.
It is considered that they probably belong to the jaw of the same individual.
M 536 (plate 31(¢), (4), (s))
An isolated left upper deciduous third molar, the posterior buccal root
of which is broken off at the base, while the other roots are broken about half-
way down. The tooth is in fairly advanced wear.
M 263 (plates 32(g); 33(d); 34(d))
This is a fragment of the right upper jaw containing DM’, DM? and M1.
The teeth are in early stage of wear, DM? being in a more advanced stage than
M?! whose posterior part of the posterior pillar is just commencing wear.
In spite of their similar appearance and their similar degree of wear, it is
unlikely that 944 and 536 (left DM? and DM respectively) on the one hand,
and 646 and 263 (right DM? and DM? respectively) on the other hand belong
to a single individual. The Dental Index has been calculated (see Section I,
chapter 5) for both pairs of teeth, and compared with the corresponding range
of variation for the extant G. camelopardalis. As seen in the following table, all
three ratios (length, breadth and Tr./A—P index ratio) fall outside the respective
ranges for the extant material:
Dental index Tr.|A-—P
Length index Breadth index index
944/536 .. Se Me es - 83°3 5 hig | 85°7
646/263... AS ea ae . gg'0 74:8 75°7
Range of DM?/DM3 in G. camelopardalis 62-7-81°9 75°5-92°3 94°0-127°4
TABLE 30
M 533 (plate 31(@), (/), (¢))
It is an isolated right upper deciduous fourth molar, with the roots broken
off at the base. The anterior pillar has just commenced wear.
M 535 (plate 31(e), (m), (w))
It is an isolated left upper deciduous fourth molar, with the anterior buccal
root broken off at the base, while the two other roots are broken about half-way
down.
M 1115 (plate 31(f), (n), (2)
It is an isolated left upper deciduous fourth molar, whose roots are broken
off near the base. It is in an early stage of wear.
(b) Adult teeth
M 532 (plates 33( f); 35(@), (¢))
This is a left upper premolar, P?, in early wear.
Buccal surface. It has the typical appearance of a paracone of an upper
premolar. Although there is a vertical furrow on the buccal surface between
460 _ ANNALS OF THE SOUTH AFRICAN MUSEUM
paracone and metacone, there is no separation on their lingual surface. Between
the apex of the paracone and the parastyle, there is a smal] enamel tubercle.
The central pit is irregularly V-shaped being open anteriorly, but closed by a
ridge of enamel posteriorly between the protocone and the metacone, while
the enamel] in the region of the protocone invaginates into the central pit as a
double fold.
Lingual surface. ‘The surface is fairly rugose. There is a ridge of enamel
opposite the centre of the protocone, while this ridge increases posteriorly to
form a broad entostyle.
M 531 (plates 33(2); 35(d), (#))
A left P?. This is a very worn tooth.
Be Makapansgat
maa an oy ar Rey Cadults)
p3
m1 aa
Me “ .
Me baPiS 2) ele S ine UNOS eee 8 ae
Mo e °
M3 PEPPR il emera ye Sten se
15 20 25 30 She, 40 45 mm
Fic. 20. Dimensions of adult teeth of Makapansgat Giraffa plotted against ranges of
variation in modern Giraffa camelopardalis.
—— = A-P length. ------ = Transverse,(breadth). . = Makapansgat specimens.
Lingual surface. A cingulum is present; rugose lines may be seen scattered
all over the enamel, but the surface has been smoothed. The enamel bulges
just below the cingulum and there are marked bulges at the bases of the
parastyle and the metastyle.
Occlusal surface. The centra] pit is almost obliterated, the enamel of the
two sides being in close contact or even fused, and there is a small island of
enamel cut off from the central pit in the direction of the metastyle.
The roots are very similar in shape to M 1114 (vide infra), but they are
ever shorter, their tips tapering to a sharp point very rapidly.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 461
M 264 (plates 33(2); 35(4), (F))
A right P?, which is in very advanced wear. The general appearance is
that of M 531, except that the rugosity is more widespread and that the small
island of enamel on the occlusal surface which leads up to the metastyle is in
contact with the enamel of the rest of the central pit. The parastyle is markedly
curved and twisted posteriorly on its own axis, as in M 531 and M 1114. As
in M 531 too, the entostyle is not present because the tooth is worn above the
level at which it projects in M 1114 (vide infra).
Roots. They are rather smaller than in other teeth and there appears to
be a constriction at the base of the posterior buccal root.
M 1114 (plates 33(g); 35(¢), (g))
A right upper ? third premolar, fragmented and in advanced wear, with
the metacone broken off.
Lingual surface. A cingulum is detectable and there is a rounded bulge
below it. There is a minute enamel nodule on the antero-lingual aspect in
the region of the protostyle. It is fairly rugose. There is a marked entostyle
on the posterior part of the lingual surface. Half the buccal surface is missing.
The anterior half resembles the paracone of a premolar in that the enamel
ridge passing to the apex is thick, rounded and triangular in shape and the
enamel is folded to form a prominent parastyle. The central pit is irregularly
V-shaped. It narrows towards the parastyle, but broadens out towards the
metastyle where the enamel is partly broken away. As mentioned in M 264,
M 531 and M 532, the buccal enamel of the protocone evaginates into the
central pit and with wear it isolates (M 531, M 264) an island of enamel of the
central pit in the region of the metastyle. This feature is common to the pre-
molars in Giraffa and Sivatheriinae.
Roots. ‘There are three roots. The lingual root is very broad at the base,
triangular in shape, and the two buccal roots are smaller and separated by a
V-shaped interval from each other and from the lingual root. The roots are
all short. The two buccal roots are broken off half-way.
M 552 (plate 31(g), (2), (w))
An incomplete, isolated right M1; the roots are broken off at the base.
The tooth is in quite early wear.
M 551 (plate 31(h), (£), (*))
It is an isolated left upper molar, M4, with the roots broken off at the
base. Very slight degree of wear can be observed only on the anterior cones
of the anterior pillar.
M 550 (plate 34())
This is a fragment of a maxilla containing a left M?, while the sockets of
the roots of M!—P? are visible. The tooth is very worn especially on the posterior
462 ANNALS OF THE SOUTH AFRICAN MUSEUM
aspect of the posterior pillar where the enamel has disappeared completely.
The dentine on the occlusal surface has been hollowed by an unusual type of
wear. The maxilla is rather squashed and the empty root sockets are filled with
calcite matrix. On the upper aspect of the specimen a portion of the maxillary
sinus has remained.
M 528 plates 32(/), 33(¢), 34(¢))
This is a fragment of maxilla containing left M* and M? which are in an
advanced stage of wear.
(2) LOWER DENTITION
(a) Milk teeth
M 939 (plate 37(2), (A), (-))
A fragment of a left mandible containing DM, and DMs, and part of the
socket of DM, is present.
M 540 (plate 37(4), (g), (2))
A fragment of a right mandible containing DM,—DM,.
Measurements of mandible opposite talonid of DM, (mm.)
Lingual height .. Ao
Buccal height ies ears
Thickness .. A io yO
(b) Adult teeth
M 936 (plates 32(¢); 33(4); 34(¢))
A fragmented lower right molar, probably M,.
M 942 (plates 32( /); 33(¢); 34(2))
It is the posterior pillar of a right lower second molar.
M 1113 (plates 32( 7); 33(¢); 34(4))
It is the anterior pillar of a right lower second molar. It and M 942 have
identical contact surfaces and fit each other, forming a single tooth.
M 938 (plates 32(¢); 33(4); 34(a))
It is a fragment of a mandible containing an erupting M, and there is
also a socket for the posterior buccal side of the root of a M,: the posterior root
of M 942 fits perfectly into this socket, and it is obvious that M 938 and M 942—
M 1113 belong to the same individual.
DETERMINATION OF THE SPECIES OF THE MAKAPAN GIRAFFA
It has already been stated (see introduction to the description, chapter 3)
that morphologically (non-metrically) these fossil specimens are indistinguish-
DM2
DM3
DM4
M3
40
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 463
Makapansgat
Tr: A-P Index
50 60 70 80 90 100 0 120 130 140%
Fic. 21. Transverse/A—P Index of Makapansgat Giraffa plotted against ranges of variation
in modern Giraffa camelopardalis.
@ = Makapansgat specimens.
464. ANNALS OF THE SOUTH AFRICAN MUSEUM
able from the extant Giraffa camelopardalis. Statistical analysis of the dimensions
of the specimens indicates that in general they fall well within the range of
variation of Giraffa camelopardalis (figs. 19, 20, 21, table 39). However, it must
be pointed out that in quite a number of cases, the dimensions tend to fall
towards the lower limit of and even just outside the extant range; while in
some cases, they fall beyond the upper end of this range (for instance, length
and breadth of M 646, M 944; lengths of M 938, M 942 and M 11193). This
distribution raises the possibility that the specimens may belong to another
species or subspecies.
Consequently the ¢ test was applied to both the: A-P and transverse
dimensions of each of the specimens. The results of this analysis are given in
table 31. It will be seen that the probability (P), i.e. of belonging to G. camelo-
pardalis, is less than -o1 (1%) in only five cases.
A-P Transverse
Tooth Specimen t-value P smaller t-value P smaller
than than
2
he M oat } 3°2156 ‘OI 2°1258 "05
M a f °5027 Wi "2974 8
DM* M 533 )
M 263 |
MEd j 1°358 “2 1354 “2
i Ne 535 1
939 : . 3 ,
2 M 540 -| 2663 8 1-464 2
DM M 939 |
F M 540 -$ 9029 4 9443 4
DM, M 540 6874 5 ‘6801 5
ie M 532 "3984 7 "2550 8
Pps M 1114 |
M 264 4°5118 ‘OI 1°5456 “2
M 53! |
M!? M 263
M 552 "58 6 “791 5
SDE. ,
2
” NE PBB ir NAMNGEELE 9 2°3455 “02
M? M 528 19095 “I 2°846 ‘OI
M, M 936 |
M 942 ; 3°069 ‘OI 1°074 3
M 1113 |
M,; M 938 1°64.54 ‘Ol "34.09 8
TABLE 31. Results of application of the ¢ test to the length and breadth
measurements of the teeth of Giraffa from Makapansgat. (P = probability)
A further step was to estimate the extent to which these ‘abnormal’
dimensions occur together in a single population. Because of a probable
correlation between transverse and A—P measurements in individual teeth,
the probability should only be considered for one or the other measurement,
and consequently it has been estimated separately for each dimension. In
respect of the A-P dimension, applying the binomia! distribution, the pro-
bability that in 13 cases, 4 measurements occur on a less than -o1 chance is -09,
which is not highly significant. It is obvious that P, estimated on the basis of
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 465,
the transverse dimension, is still less significant. Therefore there is no basis for
placing these specimens in a different species from G. camelopardalis.
Furthermore, because the Makapansgat teeth are mainly isolated speci-
mens and the measurements of the majority of the modern specimens were
taken on intact jaws, slight differences in the actual measurements probably
resulted. This would sufficiently account for the few isolated exceptional
results. However it has also been established in other groups of the Hopefield
fauna that the Pleistocene fossil representatives of modern species tend to have
dimensions (of teeth and skeletal parts) towards the upper end of or just outside
ranges of variation of the modern species (Ewer and Singer, 1956; Hooijer and
Singer, 1960).
B. SIVATHERIINAE
UPPER DENTITION
Milk
M 937 and M 524
For the same reasons as given for Cornelia B! and B?, these two teeth are
deciduous molars.
(2) M 937 (plate 37(¢), (¢), (J))
It is either an upper left DM? or DM?. It is unworn.
Buccal surface. ‘There is a cingulum on each of the cones which have a
typical spatulated appearance, the paracone being less curved than the meta-
cone. The general appearance is similar to the buccal surface of B? from
Cornelia, except that the central ridges leading up to the apex of the paracone
and of the metacone are much more prominent. The parastyle is broken off.
Lingual surface. ‘The crown-root junction is fragmented, but a cingulum
appears to have been present adjacent to this area, and there is a rounded
bulge below the cingulum. The enamel is fairly rugose and is thrown into
horizontal ridges on the anterior surface of the anterior cone. The lingual
surface of the protocone has a thickened pilastering which leads up to the apex.
This effect is slightly less marked on the hypocone.
Occlusal surface. The hypocone has a V-shape, while the paracone tends
to be slightly more U-shaped. The central pits are open in the region of the
mesostyle.
Roots. ‘The lingual root is broken off at the crown-root junction while
the other two roots are embedded in a fragment of the maxilla and matrix.
(>) M 524 (plate 37(¢), (), (4)
A right ? DM®.
Buccal surface. ‘There is a rounded cingulum. The metacone presents a
typical spatulated surface, with the mesostyle curving outwards and forwards.
466 ANNALS OF THE SOUTH AFRICAN MUSEUM
The paracone has a marked central costa which is very broad at the base and
which has two vertical furrows near the apex. The parastyle is prominent
with a thick rounded base. There are small nodular excrescences on the para-
style which tends to recurve near the apex to give the cone its spatulated shape.
Lingual surface. It is finely rugose. There is a rounded cingulum which is
more marked on the protocone and which has a small denticulated ridge of
enamel below it. There is a very marked protostyle on the anterior surface.
On the posterior surface of the hypocone, the enamel ridge is much less marked
and there is no cingulum on this surface. There is a small linear entostyle
present. i
Occlusal surface. ‘The central pits are wide and open, except in the region
of the parastyle where the anterior part of the protocone fuses with the paracone.
The hypocone is V-shaped while the protocone is flattened out.
Degree of wear. The anterior pillar and the anterior part of the posterior
pillar are slightly worn while the posterior part of the posterior pillar is just
beginning to show signs of wear.
M 941
This is either a right DM? or DM?+. The roots are broken away, and on
the crown only the hypocone is more or less intact. It is in early wear. This
tooth shows similar characteristics to those seen on M 937. The lingual surface
slopes at a fairly marked angle towards the apex. The buccal surface also
slopes but to a lesser degree.
LOWER DENTITION
(a) Milk teeth
M 525 (plate 32(), (4), (¢))
This is a deciduous tooth and probably a left DM, (see Dental Index of
M 525/M 539 B (DM,), vide infra). There is a small rounded cingulum which
is particularly prominent on the hypoconid. The surface of the tooth is finely
rugose and worn; there is a rather large bulge on the cingulum on the posterior
aspect of the hypoconid. There is no cingulum in the region of the entostylid
where the hypoconid fuses with the entoconid.
Buccal surface. There is a fine groove between the hypoconid and the
protoconid. The protoconid has a distinct bulge. Posteriorly the protoconid
is fused with the enamel of the entoconid, and the metaconid juts out on the
lingual aspect of this fused ridge. Anteriorly, the protoconid is continuous
with the parastylid which also juts out at right angles from the enamel ridge,
and this, in turn, fuses the protoconid with the parastylid. Consequently,
there is a V-shaped depression between parastylid and paraconid on the lingual
aspect of the tooth, although the two are fused on the lingual surface about
half-way from the occlusal edge of the tooth. There is also a U-shaped depres-
sion on the lingual side of the protoconid between the projections of paraconid
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 467
and metaconid, which are fused to each other about two-thirds of the way
down from the occlusal edge of the tooth, so that there is a V-shaped interval
_ between them on the lingual surface. There is a small depression between
metaconid and entoconid near the occlusal edge. A linear central pit stretches
between the hypoconid and the enamel ridge, which links entostylid—to—
entoconid—to—protoconid.
There is a small cingulum on the lingual surface. ‘There are two hollow
roots, a posterior one below the hypoconid and an anterior root below the
protoconid. A distinct similarity is observed between this tooth and P, of
M 553 B. The only differences in appearance being that the metaconid pro-
jects in a more lingual direction in M 525, and that the enamel ridge forming
the central pit of the paraconid in M 553 Bis circular, while in M 525 it is more
V-shaped, and that the occlusal edge of the lingual surface is not as high in
M 525.
M 539 B (plate 38)
This is a fragment of a right mandible containing a DM, which has just
erupted, and the tip of paracone of M, is visible because the surrounding
alveolus is broken away.
Buccal surface. ‘The enamel of DM, is fairly rugose. ‘There appears to have
been a cingulum and there is a large hypostylid and a broken-off ectostylid.
Lingual surface. ‘There is a rounded cingulum. Finely rugose. On the para-
conid there is a broad, flattened central costa leading up to the apex of the
pillar, while the parastylid and mesostylid are less marked; the parastylid
curves slightly inwards at its anterior extremity, so that the surface presents a
slightly spatulate appearance. The metaconid has a similar appearance, but
anteriorly it fuses with the mesostylid of the anterior cone. The lingual surface
of the anterior pillar overlaps that of the posterior pillar which in turn overlaps
that of the talonid. The talonid has a very marked central costa leading up
to its apex from the cingulum. This pillar is particularly broad at its base so
that it appears to occupy most of the surface. The anterior portion of the
surface curves slightly inwards, and comes to lie against the metastylid of the
posterior cone.
Occlusal surface. ‘The A—P axis of the lingual surface of the posterior cone
is at a slight angle to that of the talonid; the anterior portion of this cone is
angulated at an even greater angle to that of the posterior cone. The central
pits are wide and open, and on the buccal surface of the paraconid the enamel
is markedly ridged, so that it projects backwards into the central pit. The
buccal cones are V-shaped in occlusal view, the central portion of each surface
producing a ridged effect which leads up to the apex of each cone.
Roots. Through the broken mandible the top of the short central root is
visible. On X-ray (plate 38(c)) the outline of the developing tooth bud of P,
may be seen, as well as the outline of the alveolar ‘pocket’ containing M, which
is still embedded in the mandible.
468 ANNALS OF THE SOUTH AFRICAN MUSEUM
Measurements of mandible (mm.)
Breadth opposite the talonid .. 3.20
Breadth opposite M, .. Aenats ¥1-'0)
Height opposite the talonid .. ¢. 47
The Dental Index (see Section I, chapter 5) has been calculated between
left DM, (M 525) and right DM, (M 539 B), in order to determine whether
they belong to the same jaw. All three ratios fall well within the range of
variation of the corresponding index for the corresponding measurements in
extant G. camelopardalis and are very close to their mean values.
Length index Breadth index Tr.|A-—P index
M 525 /M539B ae 60-0 75°6 124°0
G. camelopardalis :
Mean oth sas 63°71 F@Qe7 123°5
Range) eae ~5 1 55:6-72"3 7029251 98°7-144
TABLE 32
(b) Adult teeth
M 1116 and M 527
These are two incisors in very early wear; M 527 is at a slightly earlier
stage than M 1116. Judging by the slope of the anterior edge of the enamel of
the teeth, it would appear that M 1116 belongs to the left side and M 527 is
right.
M 1116 (plate 36(d), (e), (f))
Buccal surface. ‘The enamel is finely rugose and there is a distinct cingulum
which bulges out on the lingual and the buccal aspect. However, the cingulum
is absent on the sides of the tooth. In buccal view, the tooth is rather V-shaped,
but the apex of the V (i.e. at the crown-root junction) is flattened, while the
upper edges of the V are slightly flanged outwards. The outer surface is
convex in a mesio-distal plane.
Lingual surface. ‘The anterior edge of the enamel is spade-shaped, while
the posterior edge is irregular ( | | ). The dentine is hollowed out, the edges of
the dentine sloping towards a deep central portion; it 1s slightly worn at its
anterior end.
Roots. The root is almost complete and rather oval in shape, being slightly
flattened anteriorly, resembling very much the root of Hopefield 4026. Viewed
from the side, the crown-root junction has an inverted V-shape.
M 527 (plate 36(a), (b), (c)) |
This appears to be a right incisor, identical in appearance and shape to
M 1116, but being a little broader from side to side and in a mesio-lingual
plane at the crown-root junction.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 469
Measurements (mm.) M 1116 M 527
Total height : oe ie ye if Ce Gees —
Crown (buccal) height St. ‘- bf ey e 37°4 38-4.
Length om ae 22°0 22-7)
Maximum height G just above the crown-root junction,
i.e. at its highest point) sb - ee Re 17°6 17°4.
TABLE 33
M 553 A (plates 39(2), (4); 40)
This is a fragment of a left mandible on which the posterior part of the
symphysis menti is present. It contains a P, about to erupt as well as the tip of
the root and I, and the breccia-filled sockets for the tips of I, and I,;. There is
no evidence of the canine. On the outer aspect, near the front of the fragment,
a large foramen mentale is present; it is oval in shape, with rounded edges,
and leading from it anteriorly there is a deep groove.
The tip of the canine is approximately opposite the mental foramen
which in turn is opposite the posterior end of the symphysis.
The superior border of the fragment is very sharp, while the inferior border
is rounded. The medial surface is smooth and convex in shape, while on the
outer surface, there is a deep horizontal furrow about one-third of the height
below the superior border. The symphysial region is markedly irregular and
its posterior end is oval in shape and flattened from above downwards.
P,
Although partially embedded in the mandible, most of the crown is visible
for description.
Buccal surface. Fairly rugose. The crown-root junction is not visible and
a cingulum formation cannot be commented on. But the hypoconid is rounded.
and bulging above the crown-root junction, and slopes gradually towards the
apex. It is split vertically by a deep groove which extends down half-way from
the occlusal surface.
On the posterior surface there is a thickened ridge of enamel, at the lingual
end of which the medial portion of the entostylid is broken off.
Lingual surface. ‘This is also fairly rugose, and its prominent feature is the
metaconid which bulges out on the surface so that there is a hollow between it
and the entoconid. But anteriorly there is only a slight depression between the
metaconid and the paraconid; the parastylid is not visible.
Occlusal surface of this unworn tooth is irregular in shape, the highest point
being the apex of the protoconid which is still fused with that of the metaconid.
The absent canine is broken off near the tip of the root. All that one can
see is its circular shape. It almost occupies the whole breadth of the bone in
this region; it is Ig mm. in diameter.
Height opposite anterior end of P, (buccal 1a) sk as = 67°7
Height opposite mental foramen se ve fe 67:2
Maximum thickness opposite P, , 26:5
Distance (direct) between anterior edge of alveolus of P, and
posterior edge of mental foramen g9°0
Distance between anterior edge of alveolus of P, “and Posterior
border of symphysis ae Se 102°5
TABLE 34. Mekal of “ane an ys
470 ANNALS OF THE SOUTH AFRICAN MUSEUM
M 553 B (plate 39(c), (d))
This is a fragment of the right half of a mandible containing P, and P,,
and showing a part of the alveolar fossa for P,. Beyond P, the sharp anterior
border continues for about 7 cm. to the broken end where there is a vertical
fracture. On the buccal aspect of the mandible there is a broad and deep
groove which almost disappears opposite P,: here the buccal surface tends to
be almost straight in a vertical direction. The Jingual surface is convex and
the inferior border is rounded, as in M 553 A.
Teh,
It is just erupting and partly broken. Part of its alveolar socket is broken
away, so that one can ascertain that it is similar in appearance to P, of M 553 A.
Ps
It is still embedded in its bony alveolar socket and only its tip is erupting.
Most of the lingual surface of the alveolus is broken off, so that a large part of
the crown is visible for study. The buccal surface is fairly rugose, while the
lingual surface is slightly smoother. |
The highest point of the tooth is the protoconid which is shaped like an
inverted V, and this cone occupies most of the buccal surface of the tooth. The
hypoconid is small and in continuity with the protoconid. Half of the lingual
surface is occupied by the entoconid and metaconid, the metaconid being
almost at a right angle to the lingual aspect of the protoconid and is fused with
it just posterior to the protoconid apex: in this way an irregular L-shaped
central pit is formed. Between the metaconid and the paraconid there is a
wide interval, the two being joined laterally by a part of the lingual aspect of
the protoconid. The metaconid meets the paraconid just above the crown-root
junction so that, viewed from the lingual aspect, there is a V-shaped interval
between them. The paraconid is angulated at right angles to the A—P axis of
the body of the protoconid, while the anterior part of the protoconid is also
angulated at right angles to the rest (body) of its own axis. Thus an oval-
shaped pit is formed between the protoconid and the parastylid. On the
lingual surface, the parastylid is seen as a prominent bulge, the apex of
which is at the occlusal junction between the recurved paraconid and the
protoconid.
Maximum breadth opposite P;_.. ae vs 35°0
Maximum breadth opposite P, is e. 27°0
Height opposite anterior end P, (buccal side) si 67°3
Height opposite anterior end P; (buccal side) .. 77°4
TABLE 35. Measurements of mandible (mm.).
M 553 B} (plate 41)
This specimen was described by Cooke and Wells (1947) as Griquathertum
cingulatum neotype.
It is a fragment of a left mandible containing M, and M,. Posterior to
M,, where M, would have been, there is a mass of limestone matrix. Anterior
6
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 471
to M, the posterior cones of P, are just erupting through the alveolar surface,
and there is a vertical fracture through the anterior cones, indicating the
early stage of eruption of P,. On X-ray, the socket for the anterior root of M,
is seen filled with breccia. The roots of the other teeth are very long, penetrating
more than two-thirds of the height of the body of the mandible. The crown-
root junction is well below the alveolar margin.
M,
The tooth is in early wear, and hypsodont in appearance. It slopes
slightly anteriorly from below upwards. The buccal surface is fairly rugose
and there is no cingulum formation visible. ‘The enamel has slight irregularities
on it, and there is a small protostylid. Looking at the posterior surface, it can
be seen that the buccal surface has quite a marked slope towards the apex, and
that, in similar fashion, the lingual surface has an even more marked slope,
especially in the region of the entoconid.
Lingual surface. The crown-root junction is not visible, but there appears
to be no, or very little, cingulum formation. The surface is fairly rugose. The
rugose lines of the enamel tend to radiate upwards from the crown-root junction
and forwards and backwards from the central costa in a fan-shaped fashion.
The entostylid is broken off, and the anterior portion of the entoconid is in
contiguity with the metastylid at the occlusal surface, but the enamel on their
lingual surfaces tends to fuse lower down. The metastylid is partly broken off
but it is fairly marked, while the parastylid is short and blade-like, and it is
sharply angulated anteriorly. The central costa passing from the crown-root
junction to the apex of the metaconid is much thicker and more marked than
that of the entoconid.
Occlusal surface. The four cones are almost completely separated from each
other. The protoconid is contiguous with the paraconid at the apex of the
parastylid, while the hypoconid meets the entoconid in the region of the ento-
stylid. But the central pits are not completely closed off where the anterior
pair of cones meets the posterior pair of cones, and the medial and lateral lips
of the central pits are rather widely separated. At this early stage of wear, the
dentine appears as a narrow strip in each cone, and the protoconid and the
hypoconid are V-shaped, while the metaconid and the entoconid tend to be
more flattened.
M,
It is in a slightly more advanced stage of wear than M,, and has the identical
appearance of M,, except that the dentine is thicker in each cone.
Height opposite anterior pillar M, (lingual aspect) 78°5
Height opposite anterior pillar M, (buccal aspect) = 75°0
Breadth opposite anterior pillar M, (maximum). . a 49°7
Breadth opposite anterior pillar M, (maximum) .. se 55°5
TABLE 36. Measurements of mandible M 553 B! (mm.).
472 ANNALS OF THE SOUTH AFRICAN MUSEUM
Note.—M 553 A and M553B must belong to opposite sides of the same
lower jaw, because of:
5
(1) the proximity of the discovery;
(2) the same dental age, judging by the stage of eruption; and.
(3) the similar size and shape of corresponding regions of their fragments and of their
teeth (P,).
That portion of M 553 B which we consider to correspond to the posterior
portion of A has been reconstructed (fig. 22). It appears obvious that M 553 A
must belong to M 553 B?; this is in accordance with the degree of eruption of
P, and the degree of wear of M, and M, in the latter fragment. Consequently
these three fragments must belong to the same individual. (See also ‘Appendix’.)
M 55381
Fic. 22. Reconstructed mandible of the Makapansgat Sivatherium olduvaiense.
M 943
This is a fragmented posterior pillar of a right M,. It has all the features
of the corresponding pillar of M, of M 553 B!, but it shows more clearly the
groove separating the hypoconid and the entoconid on the posterior surface,
just above the point where they fuse. The degree of wear and the general
features are so similar to the above corresponding specimen that they could
almost be considered to belong to the same jaw.
The lingual surface has a large piece of enamel broken off; the whole of
the buccal surface may be measured because the tooth is not embedded in its
jaw (table 40).
CHAPTER 4.
HOPEFIELD (CAPE PROVINCE, UNION OF SOUTH AFRICA)
The description of the geological features of the fossiliferous site on the
farm ‘Elandsfontein’, near Hopefield, 60 miles north-west of Cape Town
(fig. 23) has been recorded previously (Drennan, 1954; Singer, 1954; Mabbutt,
1956). All the material, referred to as ‘the Hopefield specimens’ have been
recovered from the site and are now housed in the S.A. Museum.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 473
No stratification has been found on the site which consists of approximately
2 square miles (5 km.”). “The site lies in the sand veld between the Sout River
and the Langebaan-Saldanha Lagoon. This site is 300 feet above sea-level
and is divided into wind-scoured kloofs or depressions by sand-dunes which
are either drifting or, where covered by vegetation, stationary. Ridges of
SALDANHA
Oo
BAY HOPEFIELD
A <9,
ELANDSFONTEIN Ry
,
DARLING
fe)
CAPETOWN
fe)
e)
English miles
TOMS 20)
Fic. 23. Map indicating the Elandsfontein fossil site,
near Hopefield.
ferricrete cut diagonally across the length of the site, and in places the dunes
are capped by massive calcrete mounds or flat boulders of partly silicified
surface limestone. Softer, cellular calcretes are found in certain places at the
lowest parts of the depressions. The tortuous courses of the ferricrete ridges
indicate that they are the indurated lower flanks of old sand-dunes* now
* Recent research by Mr. D. Needham, University of Cape Town, tends to contradict
that there were sand-dunes, in the geological sense, originally present. Convincing evidence
indicates that the modern ‘dunes’ are the result of a recent ‘heaping-up’ of the tertiary marine-
deposited sands.
4.74: ANNALS OF THE SOUTH AFRICAN MUSEUM
stripped bare of the sand walls. This ferruginization is usually associated with
moist ground conditions, a fairly high stable water-table and an abundance of
vegetable acids in the soil. It seems that this fossil site was at one time a large
vlei or lagoon along the edge of which animals roamed.’ (Singer and Keen,
1955):
The movement of the present sand-dunes uncover the calcareous floors
from which the fossils and artefacts are recovered.
The majority of the giraffid teeth as well as the cranial fragments have
been recovered from different localities on the ‘Main site’. However, fragments
4029, 4029 A, 4029 B and 4374 were discovered on a ferricrete deposit in
site E-extension (Tex. 1). Further tooth fragments (4030, 4030 A and 4031)
were found on and near a ferruginous plateau in ‘Buffalo Bay’, less than
50 yards from E-extension, but separated from it by a large sand-dune, and
it is quite likely that the ferricretes of the two localities are in continuity
through the sand-dune. |
As a matte of interest, specimens 4029 (heavily encrusted with ferricrete)
and specimen 4028 A (found on a calcareous floor and only slightly ferrugi-
nized) were X-rayed together at a distance of 3 feet with 115 kv. for 2 seconds.
The skiagram showed that 4029 was far denser than 4028 A, which possibly
signifies that it may be more heavily fossilized. ‘The disparity in the skiagrams
is also reflected in the Uranium (U,O,) content (by courtesy of Dr. K. P.
Oakley, British Museum of Natural History), that of 4028 being 14+2 p.p.m.
and that of 4029 being 4-+2 p.p.m. This indicates that the latter may have
come from a higher level, and that conditions for fossilization may have been
more favourable. The fluorine contents (by courtesy of Mr. H. E. Krumm of
African Metals Corporation, Bellville, Cape) are 1-983°% and 1:663°% respec-
tively, and the nitrogen contents are 0-11 and 0-085 respectively.
List OF MATERIAL
Hopefield S.A. Museum
No. No.
A. GIRAFFA 3345 11716 right P?
B. SIVATHERINAE
(1) Cranial fragments 4372B 11717 base of a skull
4372 ena] posterior horn-core
4372 A 11717 fragments belonging to 4372
4373 11718 posterior horn-core
4373 A 11718 anterior horn-core
4373 B, CG 18 yp te) fragments belonging to 4373
(2) Teeth
Upper 4025 11719 right P®
4027 11720 right M+
4023 11721 left M?
4024. 11722 left M?
Lower 4026 11723 iNCisOr |
4374 11724 left M, or M,
4028-4028 A 11725 fragment of left mandible with M,—M,
4029— 4029 B 11726 fragment of left mandible with M,—M,
4.030 119727 right M,
4.031 11728 right M,
4030 A Tali 7207 fragments of roots and pillars
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 475
DESCRIPTION
A. GIRAFFA
3345 (plate 45(c), (d), (2)
This is a right P?. The tooth is in late wear.
Buccal surface. There is a well-defined cingulum. The surface has a ‘W’
formation with marked parastyle and metastyle as well as a marked paracone.
Between the latter and the two styles there are deep grooves. In anterior or
posterior view, the lingual and buccal surfaces slope towards each other in the
direction of the apex.
Lingual surface. ‘The enamel is finely rugose and there is a well-marked
cingulum. The lingual surface slopes sharply from the cingulum towards the
apex in a buccal direction. There is a smooth surface on the posterior aspect
of the tooth indicating contact with P?.
Occlusal surface. ‘The enamel of the paracone is arc-shaped, while the
occlusal edge of the lingual surface is more or less straight. The central pit is
shallow (because of the marked attrition) and it is slightly curved, the ends
pointing towards parastyle and metastyle.
Roots. ‘There are three roots, one lingual and two buccal, the one buccal
being broken off at its base, while the other is partly broken. The roots are
very short and tend to be triangular in shape.
Diacnosis. Its features and dimensions exclude it from the Palaeotraginae, such
as Giraffokeryx and Okapia. The flat profile and slope of the lingual surface of
the tooth are uncommon features among G. camelopardalis in which a rounded
bell-shaped body is the typical shape, but they are observed in G. gracilis
(Arambourg, 1947, pl. XXII, 1a). The only premolar known to represent
G. gracilis is a P* which is smaller in its overall dimensions than that of G. camelo-
pardalis.
Although 3345 falls within the range of G. camelopardalis both for A-P
and transverse dimensions it tends to be at the lower end of the range. As it is
not possible to compare 3345 directly with a G. gracilis specimen, it is tenta- —
tively assigned to G. camelopardalis. This view is strengthened by the fact that
the Transverse/A—P index of 3345 is identical to the mean index (123) of P?
in G. camelopardalis. Nevertheless it should be kept in mind that the non-
metrical features of the Hopefield specimen are very suggestive of those of the
G. gracilis premolar. This fact may be more significant than its metrical
relationship to G. camelopardalis because of the extensive range of variation of
the latter (table 7).
B. SIVATHERIINAE
(1) CRANIAL FRAGMENTS
4372 B (plate 42(a), (4))
Five fragments, found in situ associated with the horn-cores 4372, 4373
and 4373 A, were reconstructed to form the base of a skull in the region of the
4.76 ANNALS OF THE SOUTH AFRICAN MUSEUM
foramen magnum. It contains portions of the two occipital condyles, most of
the lower and most of the upper border of the foramen magnum and a portion
of the occipital bone. On the inner aspect of the fragment, the bone structure
consists of numerous large and small inter-connecting sinuses. The medial
edges of the condyles project inwards slightly, decreasing the side-to-side
breadth of the foramen magnum. The medial condylar surfaces are not com-
plete, while the lateral articular portions are missing. The medial surfaces are
slightly convex from side to side and fairly convex from front to back. There
is a small groove on the postero-medial aspect of each: condyle where it is
continuous with the base of the skull. The striking features of the foramen
magnum, viewed from the inferior aspect, are its large size and its almost
circular appearance.
The postero-superior border of the foramen magnum forms a broad,
flattened arc rather than the acute-angled arch of the modern giraffe. Although
a portion of the bone is missing above this region, it is clear that the protu-
berant mass of solid bone where the two exoccipitals and the supraoccipital
fuse in a triradiate fashion, which is present in the modern giraffe, is absent
in the fossil specimen. In fact it appears that there is a thickened pillar of bone
(the region of the exo-supraoccipital suture junction) leading up to the occipital
crest. This feature is very similar to that observed in Szvatherium giganteum (see
Falconer and Cautley, 1846-9, plate XCII, 1c), except that in the Hopefield
specimen the postero-superior border of the foramen magnum appears broader
than in giganteum. Furthermore, the postero-superior portion of the occipital
condyle is angulated quite differently to the foramen magnum than in the
modern giraffe in that the plane of this region of the condyle is parallel to the
occipital surface, while in the modern giraffe it is almost at 90.° This may
indicate that the skull was balanced on the vertebral column at an obtuse
angle, rather than the right angle between the vertical axis of the upper
cervical vertebrae and the plane of the base of the skull of the modern giraffe.
Mechanically this would be in accordance with the short neck of the Siva-
theriinae, which probably contained a group of powerful extensor muscles to
assist In maintaining the balance of the large skull with its massive, heavy,
curving horns.
Measurements 4372 B Sivatherium giganteum*
mm. inches inches
Minimum intercondylar breadth .. 56-0 ee 2°6
Foramen magnum:
Maximum A-P, internal .. zh 51 2°0
Maximum A-P, external .. ate 58 2°3 Pe!
Maximum breadth, external Me 58 2°3
Maximum breadth, internal st 61 2°4
TABLE 37
1 Falconer and Cautley, 1836.
4372 (plate 43)
This is a Sivatherine posterior horn-core (so-called ‘antler’?) which was
found to be fairly complete after reconstruction of the fragments. The base is
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 477
pear-shaped, narrow anteriorly, the posterior part being rather globular. The
lower part of the horn-core, as high as the flange, is hollow, narrowing
superiorly, and only at the lowest end (for c. 12 cm. from the broken edge) is
sinus formation visible. The base is fragmented and incomplete, the anterior
portion missing and the flange broken off at the anterior border. The body of
the horn-core is fairly complete, but the tip is missing.
Just above the base the core turns outwards, and at the base of the second
knob it begins to twist posteriorly and laterally, so that the outer surface tends
to face anteriorly. The posterior border is rounded; the anterior border at,
and above, the flange narrows (and it is more angular than at the base) while
nearer the tip, above the second knob, the anterior border becomes rounder.
There are deep longitudinal grooves running up on the medial convex
surface. There are three grooves very close together at the base of the flange
apparently diverging up from a single point. At the upper end of the flange
the posterior groove swings sharply towards the posterior border, while the
anterior two slowly separate, and the posterior of these two swings more acutely
backwards opposite the second knob. The anterior groove runs up parallel
to the anterior border as far as the second knob where it crosses the medial
surface posteriorly and upwards towards the tip. In addition, arising from the
anterior part of the base, there are three other grooves which travel obliquely
upwards (from a point behind that of the above-mentioned grooves) towards
the posterior border opposite the middle of the flange, and there they tend to
disappear towards the back.
Measurements (mm.) :
Circumference at base .. ar c. 4.00
Circumference between flange and knob 2 ae ¢. 330
Circumference 100 mm. above knob 2 .. a, c. 265
Circumference at tip ‘ ah ag
Total length along posterior border... die c. 590
A-P Side-to-side
Base ae aP eA Gat 70 G27
Between flange and knob 2 i. ae oie ¢. 130 82:0
Above knob 2 pee mm. ) ah a Nae QI‘O 72:0
At tip = i : oe on ae = =
TABLE 38
4373 (plate 44)
A posterior horn-core, the partner of 4372. No additional features are
observed on it as it has the same general appearance of 4372. But, viewed
from the front, the anterior border has a distinct sinuous (‘twisted’) appearance
(see also Old. 86 (BK II)). It is rather incomplete, especially on the anterior
border and convex surface. Therefore, the measurements would all be approxi-
mate and as they would add nothing to what has been obtained from 4372, no
measurements are given.
4373 A (plate 42(c), (d))
This is the posterior portion of an anterior horn-core found in association
with 4372, 4373. One surface is markedly concave, leading from the base to an
478 ANNALS OF THE SOUTH AFRICAN MUSEUM
irregular rounded knob (cf. C 431 from Orange Free State). No sinuses are
visible at the base which is broken away. This is the first Sivatherine anterior
horn-core to be described from Africa. Thus it becomes the paratype for
Stvatherium olduvaiense (see ‘Discussion’, Section III).
Height of fragment .. 160 mm.
Tip of fragment .: 55 mm. (transverse) X 59 mm. (A-P).
4373 B
A number of small fragments found with 4373 which cannot be fitted into
the reconstruction.
4372 a, b, c, etc.
Groups of fragments found in association with 4372. Some are tiny
fragments of the horn-core, but there are also a few fragments of skull, too
small for identification or description.
(2) TEETH
4025 (plates 45(2), (/), (g); 50(2))
This is an isolated tooth, a right P3.
Buccal surface. There is a rounded cingulum. The posterior half of this
surface is missing. The parastyle bulges markedly below the cingulum, while
a deep cleft separates the parastyle from the paracone.
Lingual surface. ‘The enamel is fairly rugose and there is a marked rounded
cingulum.
Roots. There are three roots. The lingual root which represents the root
of the protocone is massive and rounded, and it projects medially. The buccal
roots are broken off, but the anterior appears larger than the posterior root.
General shape. The tooth is very large, the buccal surface being flattened,
while the lingual surface is arc-shaped. The central pit tends to be U-shaped,
its enamel surfaces being widely separated in the centre; the ‘arms’ of the U
taper towards the anterior and posterior ends of the buccal surface. The
enamel on the lingual aspect of the pit is evaginated into it posteriorly, and the
dentine here contains a rounded cone of enamel (cf. Makapansgat, M 1114).
The paracone is wide from side to side, and its dentine tapers both anteriorly
and posteriorly.
4027 (plates 46(a); 47(a); 48(a)).
An incomplete isolated right upper molar, probably M1}, in rather advanced
wear.
Buccal surface. Marked, rounded cingulum. The tooth is rather fragmented
and the paracone is missing. The metacone is similar to that of M?, 4023 (vide
infra), and the mesostyle again is prominent.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 479
Lingual surface. The rugosity is finer than in the other teeth. The cingulum
is marked and rounded. There is a bulge below the cingulum which is more
marked in the anterior than in the posterior pillar. In the posterior pillar, this
bulge gives way to a sharp outward angulation of the lingual surface in the
direction of the apex, similar to 4024 (vide infra). The two pillars meet on the
lingual aspect, similarly to those of 4024, 1.e. the folding of the two contiguous
enamel sides of the pillars being in an anterior direction.
Occlusal surface. The dentine is hollowed out on the 4 cones, forming an
arc in a side-to-side direction. The central pits tend to be U-shaped: as in
many of the other upper molars, the enamel on the lingual side tends to be U-
shaped, while on the buccal side it is V-shaped. The U-shaped side however
is more worn than the V-shaped one.
The inner arms of the two pits are not continuous, although the enamel
of the two arms are touching each other. As in M? (4023), the posterior part
of the pit of the anterior pillar has an evagination directed buccally towards
the mesostyle.
Roots. ‘The lingual root is broad, thickened above each pillar and a vertical
groove on the medial aspect demarcates the two parts. The posterior buccal
root tends to be oval in shape and flattened from front to back in an antero-
posterior direction. The anterior buccal root is missing.
4023 (plates 46(d); 47(a); 48(a))
An isolated left M?, found in association with 4024, with which it estab-
lishes a contact. The tooth is in a rather advanced stage of wear.
Buccal surface. ‘This is similar to 4024 (vide infra) but the mesostyle, which
is here complete, is very marked and appears to be associated with the posterior
rather than with the anterior pillar. In the central hollow of each pillar there
is a slight ridge of enamel (the costa), stretching up from the cingulum towards
the apex of the tooth. This is more marked on the anterior pillar. The para-
style is broken off.
Lingual surface. There is a marked cingulum below which there is a slight
bulge in each pillar. The enamel is rugose. The enamel is raised in a marked
ridge on the anterior aspect of the tooth (protostyle).
Occlusal surface. ‘The anterior pillar is again larger than the posterior one,
and both tend to be arc-shaped on the lingual aspect. The slope of the buccal
and lingual aspects are similar to those of 4024 in that the lingual surface of the
protocone has a more vertical slope than that of the hypocone, while this is
reversed on the buccal surface where the metacone is more vertical than the
paracone. The central pits are more V-shaped, although the apex of the V,
which points lingually, is slightly flattened. The inner arms of the V of each
pillar become continuous with each other in the region of the mesostyle. Just
on the anterior side of this point the pit of the anterior pillar has a marked
evagination, which almost reaches the enamel junction of the two pillars on
the lingual aspect.
480 ANNALS OF THE SOUTH AFRICAN MUSEUM
Roots. ‘They present an identical appearance to those of M? (4024), but
the tip of the lingual root tends to be angulated in a vertical direction, and the
lingual root is shorter than the buccal ones. The posterior buccal root is
larger than the anterior one; both are triangular in shape. The tips of
the anterior and posterior buccal roots curve anteriorly and posteriorly
respectively.
4024 (plates 46(c); 47(¢); 48(¢))
An isolated left M?. It is fairly complete, rather cracked, with most of the
buccal roots missing.
Buccal surface. The enamel is rather rugose and not as smooth as in the
lower teeth. The cingulum is well marked and rounded but in the centre of
each pillar the area just below the cingulum tends to be scooped out. The
cingulum becomes continuous with parastyle, mesostyle and metastyle, forming
costae, so that viewed occlusally, the surface has a hollowed-out effect between
these three costae. )
Lingual surface. ‘The enamel is rather rugose. Below the rounded cingulum
there is a slight bulge which is more emphasized on the posterior pillar. The
enamel is thrown into horizontal ridges on the anterior and posterior surfaces
about 5 mm. from the crown-root junction (forming the protostyle and hypo-
style respectively). ‘The lingual and buccal surfaces of the tooth tend to slope
towards each other in the direction of the apex, as in 4023. On the posterior
pillar, there is a marked depression between the hypocone and the metastyle,
which tends to accentuate the metastyle. Similarly, there is a depression
between paracone and parastyle. This ‘pinched’ effect emphasizing the meta-
style and parastyle is found close to the crown-root junctions and is a common
feature in Sivatherines (e.g. F 3655, C 426, MMK 3685). This effect may
also be seen on the posterior side of the posterior buccal] root (e.g. C 426,
4024). The junction between the anterior and posterior pillars is not marked,
the enamel surface of each pillar being contiguous for about 6 mm. towards
the centre of the tooth.
Occlusal surface. The shape of the lingual enamel edge of the two pillars
is V-shaped, but the posterior is much narrower and smaller, the latter effect
being produced by the more acute slope of its lingual surface towards the apex.
The central pits are U-shaped, the central portion of the U being fairly wide,
while the enamel sides of the ‘arms’ tend to come close together. The outer
arms of each pit curve, one towards the parastyle and the other towards the
metastyle, while the two inner arms are directed towards the mesostyle. The
mesostyle is missing in part. The central portion of the U in the anterior pillar
has a slight evagination posteriorly in a lingual direction. ‘The image of this
in the central pit of the posterior pillar is broken.
The tooth (as well as 4023) illustrates very well the typical occlusal wear.
The enamel and the dentine are raised in a straight line from side to side in
the centre of each pillar, while they are well worn along the anterior aspect
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 481
of the anterior pillar and the posterior aspect of the posterior pillar. Maximum
wear has occurred along the contiguous surfaces of the two pillars.
Roots: The lingual roots of each pillar are fused by a plate, so that in effect
one only finds one broad root on the lingual side, with the anterior pillar
reflecting the largest bulge on the root. The two buccal roots are separated,
but the posterior one is broken off near the base and the anterior root is broken
at the crown-root junction. The lingual root is angulated in a medial direction.
The buccal roots are arranged so that the posterior root points backwards, and
the anterior one forwards and outwards.
4026 (plates 45(4), (¢), (7); 50(4))
This is an incomplete isolated incisor. Assuming for the purpose of
description that it is a right incisor, the anterior half of the crown is broken,
mainly on the buccal aspect. The enamel is finely rugose. The enamel on the
occlusal edge of the tooth on the buccal aspect is broader than it is at the
crown-root junction. Looked at from the side, the ‘apron’ appearance of the
enamel is again observed. On the posterior surface the enamel has a sharp ridge
running up from the cingulum to the occlusal surface. The posterior surface
on the lingual side of this ridge has no cingulum. Near the occlusal edge this
ridge is worn, suggesting contact with a contiguous tooth. The lingual aspect
of the tooth has no enamel, indicating that the tooth is in advanced wear, and
the dentine is slightly raised at the crown-root junction.
On the buccal aspect, in the centre of the tooth, the enamel presents a
groove which extends from the occlusal edge to the bulge above the
cingulum.
Measurements of 4026 (mm.)
Total height .. en A a ae ies 72+
Crown (buccal) height a a es Se BT)
Length (A-P) .. ey ee: Be a Soha
Maximum breadth at the crown-root junction .. 18-4
4374 (plates 46(d); 47(¢); 48(d))
An isolated, very fragmented lower left molar, M, or M,, found near
4029, 4029 A and B. It consists of the posterior root, part of the hypoconid
and entoconid and a small piece of the protoconid. The tooth is very worn.
The buccal enamel is rather rugose, the region of the cingulum is broken away
and the lingual surface is also broken. In occlusal aspect, the central pit is
visible, rather flattened in shape, with its anterior portion curving towards the
metastylid. :
Measurements of 4374 (mm.)
Buccal height a eo Pe oy
Length of the hypoconid .. c. 28
482 ANNALS OF THE SOUTH AFRICAN MUSEUM
4028 (plates 46(e); 47(e); 48(e))
A fragment of left mandible, containing M, and M, (4028 A), in advanced
wear.
Ms;
Buccal aspect. ‘The tooth is hypsodont and the enamel is fairly rugose.
The cingulum is marked, more particularly on the posterior pillar and on the
talonid; it is more rounded on the buccal than on the lingual side where it
is only represented by a faint ridge. At the anterior part of the posterior pillar,
the cingulum bulges in the region of the ectostylid.
Similarly on the talonid, at the anterior end of the cingulum, there is a
nodular hypostylid. The protoconid tends to be angular on the buccal side,
and even presents a slight vertical central ridge, which is not observed on the
hypoconid.
Lingual surface. ‘The rugosity is smoothed and even disappears opposite the
centre of each pillar where there is a slight bulge (costa). At the base of the
crown there is a bulge on each pillar, the posterior pillar having the larger bulge.
The bulge on the posterior pillar, which is unusually prominent and extensive,
tends to resolve itself in an upward direction in three ill-defined ridges: the
posterior ridge travelling in the direction of the entostylid, the central ridge
(costa) travelling towards the apex of the tooth, and the anterior ridge in the
direction of the metastylid. The enamel of the crown of the anterior pillar and
of the talonid is broken off.
About 30 mm. above the cingulum of the posterior pillar, on the lingual
surface, the enamel forms a horizontal ridge which extends across the anterior
pillar and towards the top of the talonid. There are also a few other irregularities
of the enamel giving the appearance of ridges.
The buccal surface of each pillar looked at from the side is vertical, whereas
the lingual surface is slightly angulated laterally towards the apex.
Occlusal surface. The protoconid and hypoconid form a triangle, but the
apex on the buccal side is an obtuse angle of about 110°. The talonid is large;
its occlusal view presents an oval surface, flattened from side to side, and its
A-P axis is at an angle of 45° outwards to that of the other two pillars.
The central pit is irregular and flattened from side to side so that in the
centre of each pillar the enamel of the two sides of the pits is almost touching.
The pit is confined to two pillars and only extends very slightly on the occlusal
surface into the talonid. The enamel on the buccal side is fairly regular and
wavy, being indented at the junction between the two pillars, whereas the
enamel at the lingual side gets broader and evaginates towards the parastylid
and also towards the metastylid.
Roots. There are two roots: the anterior pillar having a single root which
tends to bulge on both the buccal and lingual sides, these bulges being connected
by a short flattened plate. The root tends to be vertical and it is separated by
a triangular interval from the root of the posterior pillar.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 483
In the region of the talonid the root is also thickened and continuous with
the thickened portion of the posterior pillar by means of a hollowed-out plate,
so that in effect one observes one massive root. The tips of the roots tend to
curve buccally.
M,
Buccal surface. ‘There is a cingulum which is not as rounded as in Mg, but
on the lingual surface it is slightly more exaggerated than in M;. Also on the
anterior surface of the anterior pillar, the cingulum is well marked. In the
region of the ectostylid the cingulum tends to bulge somewhat.
Lingual surface. There is a slight bulge above the base of the posterior
pillar, and the resolving upward ridges are similar to those in Mg, but less
distinct. The bulge above the cingulum of the anterior pillar is less marked,
and is really just a thickening of the cingulum; it appears to lead up to the
broken parastylid. ‘Towards the posterior part of the anterior pillar, another
ridge (costa) stretches upwards from the cingulum towards the apex of the
tooth, and between this ridge and the parastylid there is a slight depression.
The enamel forms numerous horizontal ridges.
Occlusal surface. Each pillar has a separated central pit: the pits are only
very slightly separated in the region of the metastylid; the posterior one tends
to be the more irregular, because each end of its arc sweeps towards the
metastylid and the entostylid respectively. ‘The central pit of the anterior
pillar broadens out anteriorly. The enamel surfaces of the two pillars sweep
in towards each other at a sharper angle, so that they meet in a deep wedge
between the two pillars. Once again the protoconid and hypoconid form
angular apices on the buccal side; the apex of the posterior pillar being slightly
more rounded.
On the occlusal surface of the posterior pillar, the dentine has a shallow
pit on the hypoconid. This is probably a post-fossilization artifact.
Looking from the anterior aspect, the crown-root junction presents a
horizontal line from the lingual surface towards the buccal, and just near the
buccal surface the enamel projects down sharply like an apron. This is the
best example of this ‘apron’ effect which is, in general, better displayed in the
lower than in the upper teeth.
Roots. The posterior root is broad when viewed from the posterior aspect
and towards the tip it curves posteriorly. ‘The anterior root has a similar
appearance, but the tip is shorter and less curved. On both roots, the buccal
and lingual surfaces bulge more than the central portion of the root. At the
crown-root junction, between the two pillars, an accessory rootlet projects
down from the anterior pillar for a short distance. The tip is broken.
4029 (plate 49)
Part of the horizontal ramus of a left mandible with M,, M, (4029 B)
and part of the root of M,. The teeth are in advanced wear.
484 ANNALS OF THE SOUTH AFRICAN MUSEUM
M;
Almost complete, deeply stained by ferricrete (see Introduction to
Chapter 4).
Buccal surface. It is fairly rugose with a markedly rounded cingulum. The
ectoconid is marked and pillariform although broken off just below the
occlusal level. Just above the cingulum each pillar is rather rounded. Because
of the bulge, that part of the buccal surface above it tends to slope lingually,
whereas the lingual surface appears to be at about the same angle as in 4028.
The talonid appears to be angulated to the A—P plane of the pillars, but
less than in 4028. It has a rounded appearance because of a large bulge of the
crown above the cingulum.
Lingual surface. Rather broken. The cingulum is not rounded and not as
pronounced as on the buccal side. ‘The surface presents similar features to 4028.
Occlusal surface. The central pit is very similar to 4028. Despite a greater
degree of wear, the central pits of the anterior and posterior pillars are not
confluent. The ridges of enamel of each pit lie in contiguity. The buccal
enamel of the pillars is arc-shaped rather than triangular.
Roots. Appear to be the same as in 4028.
M,
Incomplete and rather cracked, but most of the features can be recognized.
The general appearance is similar to M, of 4028.
Buccal surface. Marked, rounded cingulum with a bulge above it on each
pillar. There is an obvious ectostylid. |
Lingual surface. The crown-root junction between the pillars appears to
be more angulated upwards than in 4028.
Occlusal surface. The central pit of the anterior pillar is small and asym-
metrically placed towards the anterior end of the tooth; the central pit of the
posterior pillar is very compressed in its anterior half; the posterior half presents
a small triangular pit, and is very similar to the same region in M, of 4028.
Anteriorly the pit does not extend beyond the junction of the anterior and
posterior pillars, as it does in 4028. These differences however are probably due
to wear. The general shape in occlusal view is the same as in 4028.
Roots. Viewed from the front, the apron effect of the crown-root junction
is not as marked as in 4028.
M,
The posterior root is present. It is broad from side to side, and flattened
antero-posteriorly.
Measurements of mandible (mm.)
Breadth opposite posterior pillar of M, .. c. 37
Breadth opposite posterior pillar of M, .. c¢. 47
4030
This is a fragment of a right Mg, the roots of which are completely absent.
Most of the posterior pillar has been reconstructed, while only a small piece
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 485
of the buccal surface of the anterior pillar remains. Although the tooth was
found in a number of fragments, these have been rejoined at only the obvious
contact surfaces. Because of the angulation of the fragment, the central pit
could be wider which would give the tooth an increased breadth, but it was
decided to replace the fragments at their minimum breadth.
Although the tooth is in fairly advanced wear, it is hypsodont. From a
rounded cingulum the buccal surface slopes medially at a fairly acute angle,
and just near the occlusal edge the surface tends to be more vertical. ‘The
lingual surface slopes upwards and laterally from the base, so that the width
of the tooth at the crown-root junction is far greater than at the occlusal
edge.
Buccal surface. ‘The enamel is rather rugose and the buccal surface is
irregular. The posterior surface shows a marked indentation at the crown-
root junction. There is a small ectostylid present leading from the rather
rounded cingulum of the anterior pillar. There is a small bulge above the
cingulum of the anterior pillar.
Lingual surface. Only a small fragment of the entoconid remains, but it
can be seen from this that the central costa is flattened; there is a slight groove
separating the upper portion of the costa from the fragmented remains of the
entostylid.
Occlusal surface. A small portion of the central pit remains. It can be
ascertained that the occlusal edge of the buccal surface has a broad U-shape.
4031
This is a fragmented portion of an isolated right M, in which the talonid
is fairly complete, but only a portion of the posterior pillar remains. Most of
the posterior root is present. It is most probable that this tooth and 4030,
found near it, belong to the same jaw. The tooth appears to have been in
fairly advanced wear. It is markedly hypsodont.
Buccal surface. ‘The talonid is very large, presenting a marked cingulum
above which the crown surface is not regular. There is a broad flattened bulge
above the cingulum and an indentation (convexity towards the lingual surface)
just above this at about two-thirds of the distance up from the crown-root
junction. The enamel is rather rugose. The posterior half of the buccal surface
of the hypoconid is present and has a round cingulum; the surface of the pillar
has two indentations, one just above the cingulum and one at about two-thirds
of the height from the cingulum. These abnormalities of the surface of the tooth
appear to be a purely local phenomenon (? ecological or dietary variation),
because anomalies of the enamel are also observed on Hopefield 4028 and
4029.
Lingual surface. ‘The cingulum is represented only by a thin ridge above
which there is a bulge on the talonid, and although most of the lingual surface
of the entoconid is missing, sufficient remains to indicate that there is a rounded
bulge above the cingulum similar to that in 4028 A.
ANNALS OF THE SOUTH AFRICAN MUSEUM
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MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA A4QI
Occlusal surface. ‘The occlusal surface is extremely broken but it can be
seen that the talonid is angulated slightly to the longitudinal axis of the tooth
in a lateral direction. The talonid is large and oval.
4030 A
These are a number of small fragments found in the vicinity of 4030 and
4031, which cannot be included in their structure. They consist of the tips of
two lower molar roots and many small fragments of cones.
SECTION III
GENERAL DISCUSSION AND CONCLUSIONS
SUB-ORDER Ruminantia
FamiLy Giraffidae
SUBFAMILY Sivatheriinae
Genus SIVATHERIUM Falconer & Cautley 1835-6
(Syn. InDRATHERIUM Pilgrim, G. E., 1910. Rec. Geol. Surv. India, 40, pt. 1, 63)
Stvatherium olduvaiense Hopwood. 1934*
* Hopwood’s specific name is retained, but the masculine form olduvaiensis is here corrected
to the neuter olduvaiense, a form which Hopwood used in 1936 but dropped in 1937.
1934 Helladotherium olduvaiensis Hopwood, Ann. G Mag. Nat. Hist. (10) 17, 546.
1936 Sivatherium olduvaiense Hopwood, Ann. G Mag. Nat. Hist. (10) 17, 636.
1937 Sivatherium olduvaiensis Hopwood. Dietrich, W. O., Wissensch. Ergebnisse der Oldoway
Exped. 1913. Dr. H. Reck, edit., p. 106.
1942 Sivatherium olduvaiense Hopwood. Dietrich, W. O., Palaeontographica, 94, A: 43.
1947 Griquatherium cingulatum Haughton. Cooke, H. B. S., & Wells, L. H., S. Afr. F. Scz., 43,
232.
1948 Sivatherium olduvaiense Hopwood. Arambourg, C., Mission Scientifique de l’Omo (1932-3).
T. 1, Fasc 3, Paléontologie, Paris, Mus. d’Hist. Nat., p. 376.
1948 Libytherium maurusium Pomel. Arambourg, C., C.R.S. Soc. Géol. France, Paris, p. 178.
1953 Sivatherium sp. Dreyer, T. F., Res. Nas. Mus., Bloemfontein, J, pt. 3, p. 74.
Sivatherium olduvaiense vanhoepeni subsp. nov.
1932 Orangiatherium vanrhyni van Hoepen, Pal. Nav. Nas. Mus., 2, pt. 5, 63.
Sivatherium olduvaiense haughtoni subsp. nov.
1949 Griquatherium haughtoni Cooke, Geol. Surv. Mem. No. 35, pt. 3, 58.
Sivatherium olduvaiense subsp. indet.
Two milk molars from ? Cornelia ? Florisbad. See this paper, p. 526.
Sivatherium cingulatum (Haughton)
1922 Griquatherium cingulatum Haughton, Trans. Geol. Soc. S. Afr., 24, 11.
Genus LinyTHERIUM Pomel 1892
1892 Libytherium maurusium Pomel, C.R. Acad. Sci. (Paris), 115, 100.
492 ANNALS OF THE SOUTH AFRICAN MUSEUM
CHAPTER I
SUMMARY OF OBSERVATIONS ON AND VARIATIONS
IN SIVATHERINES
A. CRANIAL FRAGMENTS
(1) SKULL
The only fragment of a skull known from Africa is specimen 4372 B from
Hopefield. ‘The predominant features are the great circular size of the foramen
magnum, the absence of the supraoccipital bulge and the orientation of the
occipital condyles (see page 475).
(2) HORN-CORES
There are two pairs of horns, the major features of which are:
(a) Anterior horn-cores. There are two African specimens from which
conclusions may be drawn, namely, at Hopefield and at Tierfontein (O.F.S.).
The Hopefield specimen is incomplete, but the available fragment corresponds
closely to the corresponding region of the Tierfontein specimen. The con-
clusions are based on the pooling of observations on both specimens. The horn
is short, with cranial sinuses projecting slightly into the base. The horn is
flattened from side to side, presenting two surfaces and two borders. The
anterior border is rounded and has a peculiar cauliflower-like prominence
superiorly. The posterior border is also rounded medio-laterally, but from
above downwards it presents a concavity facing posteriorly. The one surface
is particularly marked by deep grooves, especially near the front. As will be
demonstrated in the discussion below, the Hopefield and Tierfontein material
are to be referred to S. olduvaiense. The Hopefield specimen 4373 A becomes the
paratype, because the authors identified it as an anterior horn-core before
studying the Tierfontein specimens. Van Hoepen (1932) mentioned only the
posterior horn-core definitively, but did not recognize the other specimen
(van Hoepen’s ‘terminal fragment’) as an anterior horn-core. He also did not
indicate that either specimen belonged to the giraffids.
(b) Posterior horn-cores. Posterior horn-cores have been recovered from
North Africa—St. Arnaud (Arambourg, 1948), Ain Hanech (near St. Arnaud)
and Garet Ichkeul (Tunisia) (Arambourg, 1949)—East Africa (Olduvai
Gorge), and South Africa (Tierfontein and Hopefield). The features common
to all of these are:
(i) Posterior horn-cores are of great length, averaging 640 mm. along the
posterior border (range 560-840), and of great size, averaging 390 mm.
circumference at the base (range 330-410 mm.) (table 41).
(ii) The base is rounded or pear-shaped and is hollowed out by the cranial
sinuses.
493
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MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
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ANNALS OF THE SOUTH AFRICAN MUSEUM
(ii1) The body of the horn-core is hollow for a varying extent.
(iv) Quite distinct from Cautley’s specimen of the “‘palmated antler’ of
S. giganteum (Falconer, 1868, I, pl. 21, fig. 3),* the African specimens
generally are rounded or oval-shaped. The African specimens present a
narrow convex anterior border and a rounded concave posterior border
which also has a spiral twist, so that the anterior border becomes superior
near the tip and the posterior becomes inferior, i.e. anti-clockwise (right
horn) or heteronym (following the terminology of Lydekker, 1913).
The horns are characterized by a flange-like projection (which varies in
extent) of the anterior border just above the base, and knob-like projections
(usually at least three) of varying size along the anterior border above
the flange.
It has been clearly demonstrated in Section 1, chapter 7, that in
the modern Giraffa camelopardalis there are tremendous sex and individual
variations within the species. It has been indicated that the previous
sub-specific ecological criteria and horn variations are not valid.
There can be no doubt that the dentitions of the Sivathertum specimens
presently found at Olduvai belong to a single species, olduvaiense, in
common with those from Hopefield, Port Allan (Tierfontein), Makapans-
gat, Omo, and Garet Ichkeul. The horn-cores assigned to this species
from these sites show a range of variation as wide as that in the modern
giraffe. There are degrees of curvature in an antero-posterior direction
and also of twisting or torsion. The torsion varies from nothing to
almost 90°. This enormous range is found at Olduvai where most of the
specimens are curved in an A-P direction with only a slight or medium
degree of lateral twist, while a few are twisted markedly so that the planes
of the surfaces alter and the horn develops an ‘antler’ effect. The twist
usually occurs just at or above the flange. Unfortunately there are no
associations with these horns to indicate the sex of the animals; possibly
the males, as in the modern giraffe, show the most marked. variations.
The few Stvatherium giganteum horn fragments found may well be at this
extreme range of (? male) variation of torsion, giving a fairly constant
‘antler-like’ appearance, as described previously.
There are also variations in the number, position and size of the
knobs. In some specimens they are evenly distributed along the anterior
border. In the horns with marked torsion one or more of the lower knobs
get shifted towards the back. ‘The posterior ‘branch’ or knob on
Sivatherium giganteum may well again be an extreme example of this. In
other specimens the knobs are unevenly distributed, and in a few they
tend to be clustered close together near the base of the horn, thus leaving
a long smooth anterior border leading up to the tip.
* However, fragment AMNH 19774 (from the Siwaliks), referred by Colbert (1935) to
? Sivatherium, is distinctly rounded and is characterized by features observed in the African
specimens.
4
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 495
There is no sex dimorphism in the dentition of the modern Giraffa
camelopardalis. ‘The teeth of Stvatherium olduvaiense from each site tend to
fall within a single wide range of variation. Based upon the above
discussion, it is unreasonable to separate the horn-cores at Olduvai into
two species merely because of the fact that some are more markedly
twisted than others. The small differences between individuals produce
a gradation where the specimens at each end of the range appear to
differ fairly widely from each other. But the same, but not correlated,
difference can be obtained for the size and the shape of knobs and the
A-P curve.
Arambourg (1948) states that Lzbythertum horn-cores differ from other
Sivatheriinae, and in particular from the genus Sivathertum, because they are
not so branched. He refers to the number and size of the knobs, and the torsion
of the horns. However, the question of the ‘branching’ has been discussed above
and it, as observed in the African specimens, does not provide a generic or
specific criterion of distinction. Furthermore, the North African (St. Arnaud)
horn-cores* do not differ essentially from those of the other African Sivatheriinae.
In actual fact, it will be demonstrated later in the discussion that for many
reasons (one feature being the horn-cores), part of the Libytherium maurusium
material must be referred to Sivatherium.
(v) On the antero-medial convex surface} characteristic and more or less
parallel grooves of varying length and depth sweep upwards from the
anterior part of the base. The significance of the grooves is not clear.
Because of the fact that they appear to radiate from a single point of
origin at the base, one gains the impression that they may be of vascular
origin (one could compare these with the appearance of the middle
meningeal vessels on the interior of a skull). This view had previously
been accepted (Abel, 1904; Colbert, 1935; Arambourg, 1948).
However, first, most of the grooves do not decrease in size propor-
tionately to their distance from the origin, which one would expect if
they were vessels.
Secondly, one could surmise that a massive horn-core would require
a large blood supply, but this argument falls away when one considers
that the small anterior horn-cores (about one-quarter of the length and
* These have been studied at the Muséum National d’Histoire Naturelle, Paris, by kind
permission of Professors Arambourg and Lehman. The descriptions are not included in this
paper as they are to be given by Professor Arambourg in a paper now in preparation. However,
he has kindly permitted us to publish the measurements and their general appearance.
+ It is extremely difficult to determine to which side a horn-core belongs because of the
absence of cranial attachment. However, following careful consideration of the Olduvai
specimens 1.53 and 2.53 and of the S. giganteum specimens (Falconer & Cautley, 1846-49;
Falconer, 1868; Abel, 1904; Colbert, 1935), it has been decided by the authors that the posterior
horns are probably directed laterally and backwards, and set at an angle of about 45° to a
vertical and horizontal plane, when looked at from the front. In this way, the surface of the
horn-core exhibiting the marked grooves becomes the antero-medial surface.
496 ANNALS OF THE SOUTH AFRICAN MUSEUM
less than one-tenth of the mass of the posterior horn-cores) present equally
large or larger grooves. Furthermore, along the line of these grooves, one
does not find particularly significant nutrient foramina. In fact the
foramina are very small and are found scattered over the whole surface
of the horn-core, without any specific relationship to the grooves.
Thirdly, it would appear to be an unnecessary hazard for Nature to
have exposed large vessels of great length in such an unprotected region
as these curved horns.
The only apparent alternatives are ligamentous or cartilaginous
attachments, and on the basis of the large grooves found in other bovids
such as Bubalus (Homotoceras) where cartilage is known to surround the
horn-core, it would appear reasonable to assume that these grooves are
mainly for cartilaginous attachment, and possibly some of the smaller
grooves (those posteriorly) may be for blood-vessels. This is contrary to
Colbert’s belief (1935) that the Giraffidae never developed a horn sheath
during their evolutionary history, and that the hairy horns of the
modern giraffe denote a primitive character that typified the various
fossil forms.
(3) MANDIBLE
Making use of Makapansgat specimens M 553A, M 553B and M 553B?,
of which A and B? belong to the left side, and B to the right side of the same
mandible (see Section II, chapter 3), the left half of the mandible was recon-
structed by using the mirror image of M 553B. This resulted in a rather
robust mandible (fig. 22) which was compared with the Hopefield fragment
4029, the Olduvai specimens 1, 6, 91, 92 and 392, the type specimen of
Libytherium maurusium (Pomel, 1892) and the North African specimen 1950—I-I
(from Garet Ichkeul, Tunisia; in the Muséum d’Histoire Naturelle, Paris).
These were further compared with the juvenile mandible M 539B from
Makapansgat, and with data on modern giraffes. From this study the following
observations and inferences were made:
(a) The total length of the jaw from the anterior alveolar border at
symphysis menti to the posterior border of the alveolar socket of M, is not much
longer in Sivatherines than in modern giraffes (5-15 % difference).
(b) However, it is in the relative proportions of various significant regions
of the mandible that Sivatheriinae differ from G. camelopardalis.
(i) The modern giraffe shows an absolute reduction of the length of the
premolar-molar series (fig. 24) (for reduction of individual tooth sizes,
see “C. Dentition’, infra). |
The length of the premolar-molar series in proportion to the length
of the jaw (as determined above) ranges from approximately 60 per cent
in the fossil Sivatherines to 40 per cent in the modern giraffe.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 497
(ii) The distance from the anterior border of P, to the posterior border of
the symphysis menti is greater in the fossil Sivatherines than in the extant
group:
SIVATHERIINAE
51.5x48.7 476 x 49.7 41.5x 50.9 467x297 >390x 44.
410 x 329
UPPER
DM 4 DM 3
LOWER
331x179
§5.0x 240
686x 336 512 x 35.7 458x336- 426x304 354x259 260x182
M 3 M 2 M 1 P 4 P3 P 2
Fic. 24. Graphic representation of the relative ranges of dimensions (A—P and breadth)
of adult and milk dentitions in the African Sivatheriinae (excluding F 39, MMK 3685).
Mean dimensions are indicated.
Comparable ranges of the modern Giraffa camelopardalis are superimposed on
those of the Sivatheriinae.
mm. Percentage of
total length
1950-1-1 (North Africa) c. 146 32%
Libytherium maurusium type ¢. 100
M 553A €. 103
G. camelopardalis eaeks: LO
(iii) The distance between the anterior border of P, and the posterior border
of the foramen mentale is much greater in the modern giraffe:
mm.
M 553A 99
Libytherium maurusium Pomel—type specimen 100
G. camelopardalis 145
498
(iv)
ANNALS OF THE SOUTH AFRICAN MUSEUM
Comparing the data of (ii) with that of (iii) it is clear that foramen
mentale ‘s considerably anterior in relation to the position of the posterior
border of the symphysis menti in the modern giraffe. In the Sivatherines,
the foramen mentale is opposite the posterior border of the symphysis.
This is confirmed by direct measurements: in Pomel’s specimen of
Libytherium, they are opposite each other; in Makapansgat 553A, the
posterior border of the foramen is c. 9 mm. posterior to symphysis menti,
while in G. camelopardalis this distance is c. 67 mm.
In the modern giraffe, the symphysis menti is considerably longer than
that reconstructed in the Makapansgat specimen 553A when the measure-
ment is taken from the position of the tip of the roots of the incisors.
The measurements are 152 and approximately 100 mm. respectively.
Following on observation (iv), if, in 553A, one considers the combination
of the thickness of the symphysis (41 mm. at the anterior broken extremity ;
compared with that of G. camelopardalis at the same point, viz. c. 15 mm.),
the shortness of the symphysis and the height of the body of the mandible
(c. 65 mm.) at the level of the root of I,, it would seem that the anterior
canine-incisor alveolar arch must have been rather cup-shaped in the
Makapansgat specimen, so that the teeth would have been embedded
more vertically in the jaw than in the modern giraffe. ‘This conclusion
is supported by the appearance of the same region in Hydaspitherium sp.
(A.M.N.H. 19684) (plate 50(e)), although it is unlikely that the
Makapansgat specimen’s teeth were as vertically embedded in the bony
alveolus as those of HAydaspitherium. The Makapansgat Sivatherium
canine-incisor row was probably in an intermediate position. Further-
more as the canine has not yet appeared and the canine-incisor arch
appears to have ‘no space’ for it, considerable lateral expansion and
increase in depth of the arch would have to take place to accommodate
all the teeth. This specimen also indicates that the canine is the last tooth
to appear in the dentition, as in the modern giraffe. (See also ‘Appendix’.)
(c) Certain features concerning the breadth and height of the Sivatherine
mandibles are also important:
(i) Breadth. The maximal breadth is found in the region between the
.
a
anterior pillar of M, and the anterior pillar of M,. Anterior to this
region, the breadth decreases in a regular progressive fashion. In the
juvenile mandible fragment M 539B, the maximal breadth is found
opposite the ‘talonid’ of DM,. This corresponds to the region opposite
P, in the adult. If one now compares this with the breadth at this point
in the adult, then it appears that a growth increase of about 30 per cent
occurs between the two phases. :
Height. The maximal height is (as expected) opposite the talonid of Ms,
and anterior to this there is a progressive decrease towards the anterior
(ii)
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 499
extremity of the jaw. Once again, if one compares the height in the
juvenile at a point analogous to that in an adult specimen, then it appears
that approximately a 100 per cent increase in growth occurs between the
two phases.
The ratio of height to breadth is expressed as the index of robusticity. This
is maximal in Sivatheriinae in the region of M,—P,, but the range varies
tremendously between different specimens (e.g. 48:3 in Libytherium
(=Stvatherium) 1950-1-1; 69:1 in Olduvai 91).
B. PostcRANIAL FRAGMENTS
Postcranial fossil giraffid remains from Africa are very scarce. All the
material studied by the authors comes from Olduvai.* Further specimens from
various sites have been described by:
(2)
(4)
(d)
Stromer (1907): a portion of the proximal extremity of a femur, a meta-
carpal, a fragment of calcaneum showing the fibular facet, and one
phalanx: from Wadi Natrun, Egypt.
Dietrich (1937): a metacarpal E. 122, from Olduvai gorge; and (1942):
from Garussi-Korongo, a metacarpal numbered 1.29, a tibia fragment, a
calcaneum, a few first phalanges, astragali numbered Vo 330.1 and
Gar.K. 1.29, and 3 cubonaviculare numbered Vo 330.
Arambourg (1947): a distal fragment of a scapula from Omo, Ethiopia.
Bate (1951): a distal end of a radius and a distal portion of a humerus,
from Abu Hugar, Sudan.
The NON-METRICAL features of the Olduvai specimens which are distinctly
different from those of extant giraffes, are:
and
the
first: in the cubonavicular, the posterior articular facet which articulates
with the metatarsal is absent;
secondly, in the metatarsal the vertical anterior groove is much deeper and
more scooped out;
thirdly, Arambourg (1947) noted that the spine of the scapula was attached
to its dorsum nearer the glenoid cavity;
fourthly, Bate (1951) stated that the breadth of the shaft of humerus and
radius relative to the distal end of those bones was much greater in
Sivatherines than in the living giraffes.
Comparisons of the MEASUREMENTS of various bones available for study,
of those in the literature, produce a number of interesting features:
1. Scapula. There is an overall increase in the size of the scapula (table 42),
maximal breadth at the base showing the most marked differences from
* See also Appendix.
500 ANNALS OF THE SOUTH AFRICAN MUSEUM
the modern Giraffa. There is also a difference in the distance between the root
of the spine of the scapula and the glenoid fossa (which has already been noted
as a non-metrical feature). It may be noted that the data for S. giganteum is
very approximate to that of S. olduvaiense from Omo.
2 K S Giraffa' G. camelopardalis?
Sous ee
Measurements! 20 S eee
Se SER Mn Er
<3 § 33 2° 5 3
SO) ie 2 fi nae <t
Maximum breadth at the base,
including coracoid process .. 150 167 192 126 6162 "Rae
Breadth at collum sd Fe) (LOO! TT 110 80 75 99 94 83
Length of glenoid fossa .. ty g8 102 110-120 88 80 103 go 82
Breadth of glenoid fossa. . aa 80 82-100 78 76 85 83 73
Origin of the spine—glenoid fossa 66 118 314. 127 “feouseuas
Breadth of the fossa sub-spinosa,
10 cm. above the glenoid fossa 32 45
TABLE 42. Scapula.
1 After Arambourg, 1947. * Musée Royal du Congo Belge, Tervuren.
2. Humerus. There is a general similarity in the total length of the humerus
of S. giganteum and Giraffa, although the breadth of the distal extremity and to
a less extent of the mid-shaft region is much greater in Sivatherium (table 43,
page 501).
3. Radius and Ulna. ‘The data on the ulna is not sufficient to draw any
conclusions (table 44).
Olduvai G. camelopardalis
116 (S.A. Mus., Cape Town)
17176
Olecranon process to articular facet, along thesuperior border 184 143
Maximum height, olecranon process. . os ae ze ARAL 102
Maximum breadth, olecranon process Me Hig ef 78 76
Maximum breadth, articular process ae a in 04 87
TABLE 44. Ulna.
In the case of the radius, although the maximal breadth of the distal
extremity in S. olduvaiense and G. camelopardalis specimens is similar, a short
distance above the distal end the breadth dimension indicates that in G. camelo-
pardalis the shaft tapers very rapidly towards the middle of the shaft, while in
S. olduvaiense the bone generally remains wide. There is very little difference in
total length between S. giganteum (B.M. 39534) and G. camelopardalis, but the
other Sivatherines appear to have a slightly shorter radius (table 45, page 502).
501
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
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506 ANNALS OF THE SOUTH AFRICAN MUSEUM
4. Metacarpal. ‘There is a marked difference in length between the African
Sivatherines and G. camelopardalis, the Sivatherines being 40-50 per cent shorter,
and this also applies to other Sivatheriinae. It is here considered that the marked
reduction in forelimb length in comparison with the modern giraffe previously
noted, is mainly accounted for by the shortness of the metacarpal
(table 46).
5. Femur. The available information indicates no marked difference
between the extant giraffe and the extinct giraffid specimens. Unfortunately
no information is available on the total length of the femur of the extinct genus
(table 47).
6. Yzbia. In only one Sivatherine (B.M. 17072) could the total length be
measured, and it is approximately 10 per cent shorter than in the modern
giraffe. No definite conclusion can be based upon this single observation.
However the A—P length and the breadth are approximately the same in the
two groups (table 48).
7. Astragalus (talus): The Sivatheriinae specimens have a much greater
proximo-distal length, although the breadth is approximately the same. This
increased length of the articular surface of the astragalus (table 49) may have
been advantageous to the extinct short-legged animal for its efficient function,
that is, to flex and to extend more powerfully for greater speed. It may also have
been advantageous for more efficient weight-bearing.
Sivatherium S Giraffa
olduvaiense Ks 8 camelopardalis
() S —_——____<_—
S a :
a 3 Se ty) E f=}
4 ; £8 Oo S c
a ™~ a eS Sl = S 2 2
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_ Lal s Q, 8 : pS) = a > (To)
: : oe) M aS S ron ees
= 38 ¢ \¢ 386 82 ee
fe) O > O TRAt oH + HOH
Maximum proximo-distal length 113 112 130 112 124 99 93
A-P maximum length, medially 73 71 63
A-P maximum length, laterally 64 63 60
Maximum breadth, proximally.. 87 86 65 86 73 81
Maximum breadth, distally .. ¢. 75 76 72
Maximum articular breadth,
proximally .. ae Mel 73
Maximum articular breadth,
distally .. ive Eng anes WG 76 gI 76
TABLE 49. Astragalus.
1 Dietrich, 1942. * Colbert, 1935. * Musée Royal du Congo Belge, Tervuren.
8. Calcaneum. No difference is noted between the extant giraffe and the
extinct forms (table 50).
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 507
=
Ss
be a =
fe) 4 ~ Cr)
= * = & S
mS or) gS
ee fe ge ee
o foo) Os 8 Sa o-s oO 3 Oo
(o) [e) ees Ss = o ons im S &
x i = = S So = 3 fe) = 5,
Skee eS Loe ee eee
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Maximum length .. ae at 216 198 183 202 221
Maximum breadth .. ot ie 69 67 72 54 64
Maximum height, A—P tuberosity .. 69 =e. 62 72 62 72
Maximum length, A—P apres the
fibular facet .. 89 gI
Body length from the superior fodder
of the astragalus facet .. Se Lot 6. L1G 119 133
Minimum body breadth .. a 39 42 31 42
A-P length of fibular facet ee its
convex portion) 39 33 40 31
Breadth of fibular facet S93 Sie 28 24 25 26
Projection of heel .. bc ie 157
TABLE 50. Calcaneum.
1 Colbert, 1935.
*Stromer, 1907: ? Libytherium discovered in Garet el Muluk; fibular articulation of a right
calcaneum.
3 Musée Royal du Congo Belge, Tervuren.
Q. Metatarsal. As in the metacarpal, there is a tremendous reduction in
the length of the metatarsal as compared with G. camelopardalis. In all other
aspects, this bone of the Sivatherines is similar to that of the modern giraffe
(table 51, page 508).
10. Other tarsal and carpal bones. From the data available no distinct
differences may be observed between Sivatheriinae and Giuraffa (tables
52, 53, 54)-
Olduvai
34]
Maximum length ne a a 75
Maximum breadth -. 3h Se 70
Maximum postero-lateral thickness .. 39
TABLE 52. Os magnum.
ANNALS OF THE SOUTH AFRICAN MUSEUM
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MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 509
ei
=
S)
H
So gery ae
s Hens ais
Eee Mi ap pen RE REEL) §
Ly eg ry eS ee | hee
2) UP BN PS HENS. (OS OS Ela an
Pi rey oN necle eete oad ee ey
SAS Sachem eee 8. 295 SS
e) O O po Po Poe MAH HNO +O
Maximum breadth cia to
side) across centre EEO) STOO, - TOO, Well, E20. T2h 7128 109 96
Maximum A-P length across
tuberosity of navicular .. 106 106 95 127 98. = 98
Maximum length of navicu-
lar articular facet for
~ cuneiform be 55 ag 50 ; 69 |
Maximum breadth idem .. 37 37 35 38 3I
Maximum length of cuboid
facet for metatarsal of 61 61 55 Ole.
Maximum breadth idem .. 45 48 42 46 38°
TABLE 53. Cubonaviculare (Scaphocuboid).
1 Dietrich, 1942. 2 Musée Royal du Congo Belge, Tervuren.
Olduvai Olduvai
105 110A
Maximum A-P length .. Me 68 59
Maximum breadth (side to side) oe 41 40
Maximum postero-lateral thickness ists 23 23
TABLE 54. Cuneiform.
C. DENTITION
A large series of teeth provides ample opportunity for determining typical
features and variations of specimens from a particular site, and also for deter-
mining comparisons between dentitions from different sites. The characteristics
of the Sivatheriinae teeth studied are:
I. NON-METRICAL FEATURES
(a) Wear. Those teeth which are either unworn or in the early stages of
wear provide evidence that the teeth of the lower jaw are hypsodont, while the
teeth up the upper jaw are, in general, at most mesodont, although F 39 is
hypsodont. Asin all other palaeontological studies on dentitions, the appearance
of the various stages of wear may be most misleading if one has not had the
unworn teeth available. In the Sivatherine dentition particularly, because of
the great breadth of the tooth near the crown-root junction and consequently
because of the varying amount of dentine present, and because of the fusion of
the cones at different stages, the diagnosis of the teeth must be approached
with caution. The large collection of Sivatherine teeth assembled from various
510 ANNALS OF THE SOUTH AFRICAN MUSEUM
sites made it possible to establish a range of the stages of wear of the upper and
lower dentition from the unworn teeth to teeth worn right down to the crown-
root junction.
(b) Slope and bulge of tooth surfaces. Another important factor which requires
careful consideration is that of the various degrees of slope of the buccal and
lingual surfaces of the teeth from the crown-root junction towards the occlusal
aspect. Furthermore, in relation to this slope, it has been observed that the
profile of the enamel surface of the tooth may be either straight or it may have
a rounded bulge in the region of the crown-root junction, quite apart from the
cingulum. These variations are not only observed in teeth from the same site,
but there may be variations in teeth of the same jaw. These features are seen
particularly on the proto-hypocone (-id) enamel surface of the teeth.
In Old. 1, for example, the profile of the buccal surface on the posterior
pillar of M, tends to be vertical and flat, while in the talonid it has a rounded
bulge. Similar rounded bulges are observed in M, and P,. Maximal bulging
on the buccal surface is observed on the molars of Old. 6 and Hopefield 4029
(M,). On the other hand, Hopefield 4028 tends to show a flattened vertical
buccal surface, while another variation of the flattened effect is observed in
M 553 Bt where the profile is flat, but it is also sloping in a lingual direction
towards the occlusal surface. The same effect is seen in Hopefield 4030. In the
opposite direction, Old. 92 shows a very slight bulge above the cingulum, while
most of the rest of the buccal surface is recurved so that a slight concavity faces
buccally.
Similar variations in the slope and bulging of the enamel on the lingual
surface are seen in the upper teeth, except that, where the surface tends to be
straight, it slopes in a buccal direction and is not vertical. For example,
~ Old. tog has a slight irregularity of the enamel just below the cingulum, and
the lingual surface slopes buccalwards towards the occlusal surface. In Hope-
field 4023 there is a slight bulge below the cingulum and the lingual surface
has a slight concavity as it slopes towards the occlusal plane. In Hopefield
4024, there is a rounded bulge below the cingulum and on the posterior pillar
the concavity is quite marked. The maximum degree of this is seen in
MMkK 3685 where, immediately below the prominent cingulum, the lingual
surface slopes at an acute angle markedly towards the occlusal surface in a
buccal direction and also has a slight concavity facing towards the lingual
side.
Specimen F 39 (of the Vaal River site) slopes at a marked angle from the
cingulum towards the apex with a slight convexity lingualwards, but just beyond
half-way towards the present occlusal edge, the surface tends to become slightly
more vertical. It has been mentioned that the entire interior of the tooth is
completely filled with breccia, and on radiographic examination vertical
fracture lines can be seen, apparently being caused and filled by compressing
breccia. This could be an explanation for some of the excessive bulging of the
upper part of the lingual surface of the tooth. However, there are no distinct
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 51!
cracks on the surface of the enamel, except for a small one on the anterior edge
of the cingulum, and for some small cracks on the occlusal surface. This explana-
tion is only a tentative one, and has not been used as a diagnostic criterion for
discussion.
(c) Central pit. ‘The variations observed in various teeth are purely due to
the different degrees of wear and should be regarded as individual variations
rather than in the light of taxonomical differences. Similar shapes and variations
have been observed in specimens from all the sites in Africa.
(d) Enamel. The characteristic features and variations of the enamel of
the Sivatherine dentition may be described as follows:
(i) Rugosity
As already mentioned, in the whole family of giraffids the enamel is rugose,
and in the Sivatherines the rugosity is as variable as in other genera, namely
from a fine pattern to a gross appearance. These variations are observed in the
same specimen or in different specimens from the same site. Old. 6 for instance
is finely rugose, while Old. 1 (from the same BK II locality) shows a coarse
rugosity. Hopefield 4027 is finely rugose, while Hopefield 4023 (of the same
jaw) is coarsely rugose. However, in the Sivatherine teeth in general, the
enamel is thrown into far more numerous, closer-packed, vertical ‘ridges’ (each
of which consists of a number of overlapping vertical spikes) than in G. camelo-
pardalis, so that they tend to have a rougher appearance of the enamel than the
modern giraffe. Even when the authors have described teeth as having a ‘fine
rugosity’, the ridges—although less prominent—were still numerous and closely
packed, quite distinct from the modern Giraffa, where they are fewer, shorter
and separated from each other. The enamel pattern of the African fossil teeth
‘is more akin to S. giganteum than the available examples of the other Sivatherii-
nae, in that in the latter the individual ‘ridges’ are less spiked.
(ii) Cingulum
A cingulum is practically always present on the buccal and lingual surfaces,
varying from a thin, linear ridge to a rolled edge. The latter is usually found,
when present, on the proto-hypocone (-id) aspect, while the former is typically
found on the other aspect of the tooth. On the anterior and posterior surfaces,
the cingulum is usually absent or deficient. Furthermore, there may or may
not be a bulge related to the cingulum. On the other hand, just above (in
lower) or just below it (in upper teeth) there may even be a concavity of the
tooth surface (vide supra, (b)).
(111) Styles
Elevations of the cingulum are commonly present in the form of inter-
pillaric styles (entostyles and ectostylids), but this varies from tooth to tooth and
jaw to jaw. The recording of the presence or absence of the styles may be
5i2 ANNALS OF THE SOUTH AFRICAN MUSEUM
difficult because of the late stage of wear. Indeed, the styles vary in that they
may or may not reach the crown-root junction. When they do not, they are
usually present near the occlusal aspect of the tooth, so that in late wear the
tooth presents the appearance of not possessing that particular style. This
inference is drawn from the presence of the style on the unworn teeth.
The median costa is better developed on the anterior than on the posterior
pillar, and in the lower teeth it seldom reaches the cingulum, occasionally
ending as a rounded bulge above the cingulum. This effect is noticed in teeth
from all the sites. The base of the metastylid usually commences about half-way
up from the crown-root junction; consequently in late wear of the lower teeth,
with the ‘absence’ of metastylid and median costa, the lingual surface of the
tooth presents a rather flattened appearance. The mesostyle is always present
as a persistent, well-marked, rounded ridge extending from the occlusal edge
to the cingulum. The parastylid is usually ridged and slanting, being con-
tinuous at its base with the cingulum. The parastyle is always rounded,
prominent and vertical, with its base continuous with the cingulum where it
forms a bulge, and its apex tapering off to a point beyond the midpoint of the
tooth. ‘The entostylid is usually poorly represented, if present. However, the
metastyle presents a fairly prominent bulge in the region of the cingulum, but
it is relatively much smaller than the mesostyle or parastyle.
In comparison with the modern (brachydont) G. camelopardalis, the African
‘Sivatheriinae show marked differences in respect of the relationship of the styles
(-ids) to each other on the buccal (upper) and lingual (lower) aspects of the
tooth. In the modern specimens, on the buccal aspect of the anterior pillars of
the upper jaw, the parastyle meets the ridge-like prominent costa at an acute
angle at its base. Posterior to the median costa the cingulum runs horizontal
to the base of the prominent mesostyle. Posterior to the mesostyle, the buccal
surface of the tooth is flattened and seldom presents a visible median costa or
a metastyle. Consequently, the appearance of the buccal aspect of the upper
tooth is that of an inverted trident.
In the lower teeth, each pillar presents a similar appearance, the anterior
overlapping the posterior, the median costa being rather rounded, not only in
an A-P convexity but also in a superior-inferior convexity, so that the occlusal
tip of the metaconid and entoconid tend to bend over in a buccal direction.
Behind each median costa there is a slight groove separating it from its respective
stylid which is continuous with it about half-way up from the crown-root
junction. Below that level, the lingual surface presents a rather continuous
smooth bulging appearance.
In the Sivatheriinae, the styles and costae of the buccal surface of the upper
teeth tend to be rather parallel to each other, the meeting of the base of the
styles with the cingulum being in a broad U-shaped fashion. In the lower
dentition, as mentioned above, the entostylids are poorly marked and the
parastylids are prominent, which is the reverse of the situation in modern
giraffes. Because the parastylid reaches the cingulum near the base of the
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 513
median costa, the lingual surface has a rather scalloped appearance and is not
unlike the general appearance of the buccal aspect of the modern upper
teeth. | |
(e) Orientation. In the Sivatheriinae, the lingual (lower) and buccal
(upper) surfaces of the pillars present a generally smoother, flatter plane than
those of the modern giraffes, because the surfaces of the Sivatherine pillars tend
to lie more parallel to the longitudinal axis of the jaws than in the modern
Giraffa, where the surfaces of successive pillars tend to overlap each other more
acutely so that each pillar has the appearance of being independent of the next.
The talonid of the lower M, tends to be set at a variable angle to the
longitudinal axis of the tooth: the axis of the talonid is nearly in line with that
of the tooth (180°) in Old. 1, 3, 92, 120, Hopefield 4029, or at an angle of
about 130° in Olduvai 6, F. 3656, Hopefield 4028A.
2. METRICAL FEATURES
A-P Transverse Index
Teg =
N M Range of N M Range of N M Range of
variation variation variation
DM? 41°0 I.) 592-0 OO:
DM+* tr 46-7 We 307 I 85:0
Ps 1 44:
M? 2 41°5 38: —45- 2 509 47- —54-8 2 122°5 121°6-123°5
M? 3. 47:6 46:3-49- 3 49°7 47:0—-54- 3 10471 Q6-1-112°4
M® Bile 595) 440-60; 5 48:7 46:0-51°9 5 ORD. ape aren
DM, De oer a) i) Byipi\
DM, I 55'0 T2450 1 ASF
Pace 3. 26-0 3 18-2 17--19°8 3 70:2 65:4—76-2
Ps 3 354 34°0-36°3 3 259 23:-29°5 Sie o2) 04:0-81-3
Py 8 42:6 37°0-47°4 7 30°4 28°3-32°4 (eet Ooo TiS
M, Tei 5 One) aka eos 9 336 27°9-39°5 Ty G2sOr 52-4897
M, 12 51:2 486-55: 10 35°7 33°2—40°2 10 70:0 62:8—76-0
M,; 9 68:6 59:0-77- 106 333°6 Ss 3I- —30°7 9 49:6 41-9-60°1
I? I Zlko~ ; 1 184 1 87:6
I? 222-39 22-0227 2 17°5 17°4-17°6 2 883 76:6-80-0 -
TABLE 55. Transverse / A—P Index in the Sivatheriinae. The data exclude the dimensions
of F.39, MMK 3685 and those teeth not definitely diagnosed as either M? or M?.
(Italicized figures indicate approximations. )
_ The absolute A-P and transverse dimensions (table 55) in the Sivatheriinae
are greatly in excess of those for modern G. camelopardalis. The calculation of
the Transverse/A—P Index in both cases produces interesting observations:
(a) The African Sivatheriinae have a constantly (except for M+) smaller
~Transverse/A—P Index than G. camelopardalis for the milk and the adult, in both
upper and lower, dentitions. This indicates that the A—P length is relatively
more reduced than the transverse breadth in the modern giraffe (fig. 24). .
514 ANNALS OF THE SOUTH AFRICAN MUSEUM
Upper Lower
Tooth Series ——_—_—__—————_
Mean Range of Mean Range of
variation variation
DM3]| Giraffa camelopardalis! 86-9 78-8— 96-0 66°6 52-6— 78-2
African Sivatheriinae? 80:2 54°1
DM4| Giraffa camelopardalis 92°3 82:0-101°0 52°3 46:2— 58-0
African Sivatheriinae 85°0 43°7
P2 Giraffa camelopardalis 122°9 104°0—-149'0 ; 87°4 56°5-124°0
Stvatherium olduvaiense*® I110°O0 100°0—-119'0 78-0
African Sivatheriinae 70°2
Sivathertum giganteum* 113°0 65°4— 76:2
P3 Giraffa camelopardalis 125°l1 101°0-157°0 91g §=-: 688-1150
Stvatherium olduvaiense 128-0 126:0-130°0 74:0 65:0— 83:0
African Sivatheriinae 737% 64-0— (61-9
Stvatherium giganteum 120°0
P4 Giraffa camelopardalis 132°9 = 114"0-158°4 89:0 63-7-106°9
Stvatherium olduvaiense 130°0 =110°0—-142°0 76:0 67:0-— 88-0
African Sivatheriinae 972°7 65°5— 77°3
Stvatherium giganteum 129°0
Mr Giraffa camelopardalis 103°9 88-0-119°4 78-6 67:9— 981
Stvatherium olduvaiense 102°0 89-0-116°0 74:0 64:0— 81°0
African Sivatheriinae 122°5 = 121°6-123°5 72°0 58:4— 89°7
Stvatherium giganteum 107°0 100°0—115'0
M2 Giraffa camelopardalis 106:8 g1‘6-128-0 77°8 69°7— 90°9
Stvatherium olduvaiense 100-0 71:0 65:0— 83°0
African Sivatheriinae 104°1 96-1-112°4 70°0 62:8— 76-0
Stvatherium giganteum 97:0 94°0—100°0 69:0 66-0— 72:0
M3 Giraffa camelopardalis 107°4 Q1°5—-122°4 61:2 47°6— 76-4
Stvatherium olduvaiense 106:0 102:0—108:0 50°0 =49°0— 51°0
African Sivatheriinae Q5°0 85°3-111'1 49:6 41°9g- 601
Stvatherium giganteum 93°0 92°0— 94°0 49°0 48-0- 51°0
TABLE 56. Tr. / A-P index of the different types of teeth in the extant Giraffa
camelopardalis, and in several series of Sivatheriinae.
1 Calculated from data from U.S.A. Museums; see Section I, chapter 5.
2 From the African Sivatheriinae presently described.
3 Calculated from data, according to Dietrich, 1937, 1942.
4 From data according to Colbert, 1935.
(b) These observations have also been made for other Sivatheriinae on
the basis of the data of Dietrich (1937, 1942) for material from Olduvai, and
of Colbert (1935) for S. giganteum from the Siwaliks. Their results fall into the
range of variation of the African Sivatherine material here described. (table 56).
However, the index for Dietrich’s and for Colbert’s M1? fall outside the range:
this peculiarity, linked with the exceptional result obtained for M?! from the
other African material, suggests that the sampling of the M! presently described
has not been representative. Table 57 illustrates, for the various collections, the
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
aD
difference of their respective index from that of G. camelopardalis: it averages
10 units in the case of the African Sivatheriinae.
Tooth
Sivatherium
— 12°9
+29
== 2:9
st I°9
—6°8
—_ I*4
UPPER
African
—6°7
ae
+18°6
= PG)
—12°4
Stvathertum
olduvaiense! Sivatheriinae* giganteum®
ae,
Tae
=39
+3°1
—9°8
— 14-4
moe
LEG
— 13°0
— 4:6
—6:8
—I11-2
Stvatherium
olduvaiense1 Sivatheriinae® giganteum®
LOWER
— T2°5
—8-6
= F°2
—18°8
—16+3
—6-6
—7°8
—11°6
African
Sivatherium
—8-8
—'2-0
TABLE 57. Absolute difference between the respective Tr. / A—P index of several
collections of Sivatheriinae and that of G. camelopardalzis.
1 Data from Dietrich, 1937, 1942.
2 African material here described.
3 Data from Colbert, 1935.
(c) The significance of the difference of means between the fossil
Sivatherines and G. camelopardalis has been statistically tested, and has proved
‘highly significant’ for DMsg, P,, Ps, Py, M,, M2, M3, M3: ‘significant’ for DM,;
‘not significant’ for DM’, DM?# and M?.
(d) A further step was to estimate the Tr./A—P Index in other fossil genera
and subfamilies, namely, Palaeotragus, Honanotherium and Orasius (data according
to Bohlin, 1926). A similar low index was obtained (see table 58). Con-
sequently it appears that the same evolutionary trend of a reduction greater for
length than for breadth has been demonstrated by different phyla in the
giraffid family.
Giraffa
Tooth camelopardalis1
DM? __ 9668 (35)
DM? 86-96 (38)
DM? g2°28 (38)
DM, = 64°44 (37)
DM, 66-58 (36)
DM, = 52°30 (37)
PA 122-86 (117)
p> 125712 (120)
Pp 132-86 (116)
M? 103°91 (138)
M? 106-79 (130)
M3 107°44 (121)
P, 87-41 (109)
P; 91-95 (124)
r. 89-02 (127)
M, 78°59 (143)
M, 77°77 (135)
M; 61-21 (121)
Palaeotragus*
70°84 (2)
78°51 (6)
go-82 (7)
63°14 (5)
66-15 (6)
60-03 (6)
100°08 (7)
107°73 (9)
124°35 (9)
108-37 (13)
10501 (13)
101-62 (12)
83°90 (5)
84°47 (9)
82-03 (10)
82°35 (4)
79°45 (13)
50°41 (9)
Honanotherium?
71-91 (4)
80°65 (2)
95°53 (2)
58-82 (1)
61-90 (1)
59°37 (1)
102°54 (2)
115°41 (3)
134°77 (4)
106-11 (4
109°38 (8)
108-39 (6)
77°77 (1)
77°59 (2)
82-97 (2)
74°28 (1)
87°09 (1)
55°23 (3)
Orasius?
80-95 (1)
97°95 (2)
109°76 (3)
101°85 (2)
97°73 (3)
91-63 (3)
70°00 (1)
77°27 (1)
65°38 (1)
69°23 (1)
52°77 (1)
Sivatheriinae®
80-2 (1)
85:0 (1)
122°5 (2)
104°1 (3)
95°0 (5)
70°2 (3)
73°1 (3)
72°7 (7)
72°0 (7)
70°0 (10)
49°6 (9)
TABLE 58. Transverse / A—P index in fossil and extant Giraffid teeth.
Between parentheses ( ) is the number of specimens from which the mean has been calculated.
1 Specimens from U.S.A. Museums (see Section 1, chapter 2).
2 Bohlin, 1926.
3 African material here described.
ANNALS OF THE SOUTH AFRICAN MUSEUM -
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518 ANNALS OF THE SOUTH AFRICAN MUSEUM
Two hypotheses may be advanced to account for the relatively increased
breadth of the modern teeth and the reduction in length of the total molar-
premolar series relative to the length of the jaw (see Section III, chapter I, A):
(1) The maximal contact wear between contiguous teeth was observed in
the fossil lower jaws, especially in M, and P,, far more than in the modern jaws
where it is only occasionally observed. It is also in the lower jaw that the
greater length reduction has occurred: this may have been an attempt to
compensate for the ‘impacted’ effect of the over-crowded teeth and in order to
accommodate them. |
(ii) ‘The fact that the teeth decreased more in length than in breadth may
be explained by the fact that there is a selective advantage in a broader grinding
surface for the side-to-side masticatory movements.
Owing to the lack of a complete maxilla, one is confronted with a problem
to which an answer cannot yet be supplied, namely, the disproportion between
the respective length reduction in the total upper and lower dentitions.
(e) Dental index: The dental index (Section I, chapter 5) is calculated to
be usually smaller in the fossil genera (Sivathertum, Honanotherium, Helladotherium,
Palaeotragus, Orasius) for length and breadth (with constant exceptions in the
dental breadth index for M1/M?, M,/M, and M,/M, (table 59). Consequently,
in the dental series, a particular posterior tooth, if compared with the tooth
immediately anterior to it, is relatively longer in the fossil genera than in the
modern giraffe material. This indicates that the reduction in the length of
individual teeth has been greater in respect of the more posterior teeth.
CHAPTER 2
DIAGNOSIS OF AFRICAN FOSSIL GIRAFFID GENERA
AND SPECIES
Colbert (1935) has modified the classification of the family Giraffidae of
Pilgrim (1911), Bohlin (1926) and Matthew (1929) along sound lines. Because
of the limitation of the African material, it is not the purpose of this paper to
criticize this classification which is generally acceptable as a basis of discussion:
GIRAFFIDAE
Large, ruminating artiodactyls, with heavy, rugose cheek teeth. The skull
may or may not have horn-cores, but if they are present they show a great
variety of development. Bones of cranial roof pneumatic. Lateral metapodials
and digits atrophied.
Palaeotraginae
Primitive, medium-sized giraffids, having as a rule one pair of supra-
orbital, frontal horn-cores. There may be a second pair of horn-cores at the
9
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 519
anterior extremities of the frontal. Horn-cores in the form of simple tines, well
developed in the males, feebly developed or absent in the females. Skull usually
elongated, dolichocephalic.
Cheek teeth brachydont, with moderately coarse sculpture on the enamel.
Neck and limbs slightly elongated.
Genera: Palaecotragus Achtiaria (syn. with Palaeotragus).
Giraffokeryx
Okapia
Samotherium Alcicephalus, Chersonotherium, Shanshitherium (syn.
with Samotherium).
ae are thor doubtful status; placed here provisionally.
Guiraffinae
Large giraffids, with a moderately brachycephalic skull. Horns variously
developed, being on the parietals and frontals, and in Giraffa a single median
horn is also present, located on the nasals. Horn-cores rounded or flattened on
the ends and covered with hair. Skull roof with highly developed sinus cavities.
Cheek teeth brachydont, with heavily rugose enamel. Limbs and neck
greatly elongated.
Genera: Guraffa
Orasius
Honanotherium
Stvatherinae
Gigantic giraffids, with large, heavy, brachycephalic skulls. Horns
variously developed, being of frontal and parietal origin. Skull roof with large
sinus cavities.
Cheek teeth moderately hypsodont, with heavily rugose enamel. Limbs
not elongated but very heavy. Body heavy.
Genera: Stvatherium. Indratherium (syn. with Sivatherium)
Bramatherium
Aydaspitherium
Helladotherium
Vishnutherium
Griquatherium
Libytherium
On the basis of the above definitions, the Hopefield giraffids which form a
homogeneous group (except for one tooth, 3345, which has already been referred
to G. ?camelopardalis, see Section II, chapter 4) may be assigned to the Siva-
theriinae. Although descriptions and diagnosis of the different genera of
Sivatheriinae have been published, no diagnosis (per se) is available in the
literature for Vishnutherium, Griquatherium or Libytherium, yet the features of the
generic types are briefly commented on. Helladotherium is hornless and may be
excluded from consideration of the Hopefield material.
520 ANNALS OF THE SOUTH AFRICAN MUSEUM
Colbert (1935) proposes the diagnoses for Sivatherium, Bramatherium and
Hydaspitherium as follows:
Sivatherium
A gigantic Pleistocene giraffid, with four horns in the male, an anterior
conical pair, arising from the frontals, and a posterior, palmate pair situated on
the parietals. As in the other gigantic Siwalik giraffids there are deep pits in
the temporal fossa for the temporal muscles, and on the supraoccipital for the
neck muscles. The face is very short, the nasals being retracted and strongly
curved. The teeth are large, with rugose enamel. Body and limbs heavy, limbs
not elongated.
Bramatherium
A gigantic Upper Tertiary giraffe having four horns, two of which grow
up from the fronto-parietal region, and two of which extend laterally from the
parietals. Face short, with nasals considerably retracted. A large groove
occupies the parietal region just below the horn-core bases as an accommodation
for the temporalis muscles. Deep pits are located in the supraoccipital for the
heavy neck muscles. Teeth large and heavy, with rugose enamel. Limbs and
body presumably heavy and massive.
Aydaspitherium
A gigantic giraffid with two horn-cores, fused at their bases into one solid
mass, on the frontal-parietal region. The face is short, the nasals retracted.
There is a large parietal or temporal groove below the horn-core for the
accommodation of the temporal muscles. Teeth large, quadrate, with rugose
enamel. Limbs massive and not extraordinarily elongated.
From the above diagnoses it can be seen that these forms overlap con-
siderably, except in respect of their horns. ‘The brief descriptions, however, are
not found sufficient for positive determination of the Hopefield material, and
consequently various other details of the material were compared with other
collections, and it became obvious that the Hopefield specimens had greatest
affinity to Sivathertum olduvaiense, to which the homogeneous Hopefield group is
assigned without hesitation.
It has already been stated (Section II, chapter I) how Hopwood (1934)
first diagnosed his original Olduvai material as Helladotherium olduvaiensis on the
basis of some teeth and a partial hind-limb. Later (Hopwood, 1936), with the
discovery of ‘palmated antlers’, the material was referred to Stvatherium:
Hopwood stated however that ‘the antlers are not so widely palmated as in
S. giganteum, and terminate in a recurved point’; hence, the specific determina-
tion of S. olduvaiense.
It is now necessary to discuss and compare the material from the various
sites in Africa in regard to this diagnosis.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 521
I. OLDUVAI AND OTHER EAST AFRICAN SITES
All the specimens from Olduvai form a homogeneous group, both from a
non-metrical and metrical point of view. As a result of the fact that Orangia-
therium vanrhyni, which was discovered and actually named earlier than
Sivatherium olduvaiense, has now been invalidated and referred to a subspecies of
S. olduvaiense (vide infra), the question of generic or specific priority does not
arise any more. Consequently all the Olduvai material and the specimens
related to it are assigned to S. olduvaiense. To this group are also referred the
two specimens marked ‘Marsabit Road’, from Kenya.
Il. LIBYTHERIUM MAURUSIUM POMEL, 1892
On the basis of a fragmented mandible containing M,—P, and a part of
P, (plate 53), Pomel proclaimed a new genus and species which he envisaged
to be within the Sivatheriinae, but he does not state clearly his reasons for
creating the new genus. In 1947, Arambourg assigned giraffid teeth and a
scapula from Omo to Sivatherium olduvaiense, and in re-discussing Pomel’s
material, he gave as reasons for separating Libytherium from Siwvatherium:
(1) the premolar series is reduced; (2) P, has an open inner wall where para-
conid and metaconid remain separated; (3) the parastylid is very developed
in P, and in the three molars.
Furthermore, in a right mandible (1950-1-1, Muséum d’Histoire
Naturelle, Paris) from Garet Ichkeul, St. Arnaud (Tunisia) (plate 52(a)),
diagnosed as Libytherium maurusium, the above features are absent, and in all
_ respects this mandible is identical to specimens at a similar stage of wear from
Olduvai (Old. 6, for instance).
Arambourg (1948(a)) stated that excavations at St. Arnaud provided new
fragments of Libytherium, and in particular characteristic ‘antlers’ (“ramures’) of
the Sivatheriinae. Comparison of the non-metrical and metrical features of
these horn-cores (1948-1-2, 1948-1-1) (plate 51), as well as of a third
(unnumbered) specimen (plate 52(b)) (the cast of which has been made
available by the kindness of Professor C. Arambourg and Professor J. P. Lehman)
with posterior horn-cores from Olduvai, Hopefield and Tierfontein, indicates a
distinct similarity of all the specimens and it is considered that they must be
referred to Sivatherium olduvaiense.
However, on the basis of Arambourg’s criteria for the Libytherium maurusium
dentition, there is no doubt that the specimen on which he based these facts is
decidedly different, not only from Sivatherium olduvaiense, but also from specimen
1950-I—1 from Garet Ichkeul. Furthermore, these criteria are so distinctive,
especially the primitive nature of P,, that it is necessary to divide the North
African material into Libytherium maurusium for the type specimen, and Siva-
thertum olduvaiense for the horn-cores, mandibles and other similar specimens
(e.g. specimens 1948-1-1, 1948-1-2, 1949-2-937, 1949-2-938, 1949-2-725,
1931-45, 1931-8, 1931-8-110, 1950—1—90, 1950—-1-1 in the Muséum d’Histoire
522 ANNALS OF THE SOUTH AFRICAN MUSEUM
Naturelle, Paris) mostly as yet unpublished but examined by one of us, and to
be published shortly by Professor C. Arambourg. However, it may yet become
necessary to equate Libythertum with the rather widely varying genus Sivatherium.
Ill. GRIQUATHERIUM
Three specimens have been assigned to this genus: MMK 3685 and
M 553B? are described as G. cingulatum, and F.39 as G. haughtoni.
(a) ‘The Makapansgat specimen M 553 B* has been referred to G. cingulatum
by Cooke and Wells (1947). They state that from an examination ‘it would
appear that [these] lower teeth are of the correct size and form to belong to
Haughton’s species though probably to a smaller individual’.
From the range of variation obtained by the authors (see tables 39, 40)
it is quite clear that the Makapan teeth fall within the range of the East African
and the Hopefield material. In fact the breadth of M, of M 553 B! is the
smallest in the range. The general form and character, which Cooke and
Wells considered to be distinctive of the specimen, are identical to Sivatherium
specimens at a similar stage of wear, e.g. Olduvai, specimen 92. Consequently,
there is no basis, either metrical or non-metrical, for separating M 553 Bt
from Sivatherium olduvaiense; this conclusion is further confirmed by the other
Sivatherine material from Makapansgat described above, which are also
referred to this genus and species.
Although the lingual surface of P, of M 553 B is ‘open’ as in Pomel’s P,
(Libythertum maurusium, type), there is still a considerable amount of the crown
hidden by the bone, the tooth having just started to erupt. Because this ‘open’
appearance is confined to the upper portion of the tooth, it is estimated that it
would resemble Old. 6 for instance, at an equivalent stage of wear. Further-
more, the teeth are much larger than those of the Pomel specimen, and are
identical to S. olduvaiense. ‘The former point mentioned was the only possible
doubt the authors had concerning the determination of the Makapan M 553B
specimens as S. olduvatense.
(b) MMK 3685 (G. cingulatum Haughton 1922): ‘The designation of this
tooth presents a number of problems. From the description of this specimen
(vide Section II, chapter 2), the following characteristics appear to differ from
those of other African specimens: (1) great breadth of the tooth; (2) relative
difference in breadth of the two pillars; (3) marked cingulum forming an
unusually developed proto- and hypostyle; (4) prominence of the meso- and
parastyle; (5) the acute angulation of the lingual surface (in profile) to the
crown-root junction; (6) the abnormal length of the posterior pillar relative
to the anterior. Some, if not all, of these features may be found individually in
single specimens, often in a modified form. But the above features appearing
together produce this peculiarly gross form. It is certainly a Sivatherine, and
it has been compared to both Hydaspitherium and Sivatherium, and it has been
suggested that it is possibly even nearer Hydaspitherium (Bohlin, 1926; Colbert,
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 523
1935; Cooke, 1949; Singer, 1954). Nevertheless, those features in common
with Hydaspitherium are also exaggerated and it is more massive. On this basis,
and because Hydaspitherium has not yet been discovered in Africa, it is considered
reasonable to refer this single, very worn tooth to Szvatherium with which it
essentially shares most features.
As regards the specific designation, although it bears a resemblance to
F 2993, Old. 4 and tog from Olduvai, the latter specimens are closer to
Hopefield 4027, 4023 and 4024 and share the same range of variation.
MMK 3685 tends to fall just outside this group, but it is a single specimen
and no other material from Africa can yet be referred to it. Furthermore there
are no associated skeletal remains of the same animal. It is probable that if
Cooke and Wells had all the comparative materia] from Olduvai and Hopefield
available in 1947, they would not have referred M 533B! to G. cingulatum. It
could almost be considered as a nomen vanum, but in the above discussion its
generic nature is established. Consequently in the present state of our informa-
tion it would be preferable to retain cingulatum as a species of Sivatherium.
Fic. 25. Superimposition of dio-
ptographs of the lateral aspects
of F.39 ( Ver2Q80r Cite toe Ne
and Hopefield 4025 (------ 22 ibhe
probable crown-root junction of
F.39 is superimposed on that of
the other two premolars. The
crown of F 2980 is worn right
down to the root and beyond the
crown-root junction except for a
minute fragment buccally (on the
left). The buccal border of 4025
is broken away so that the overlap
of F.39 is only apparent.
1 Crown-root
t: fe ce ogee
bs aN o e e
Levi Me JUNCTION
Nz N \
: Se \;
q: 2 : al Se \
“, areas are a
re °° peo f
(c) F 39 (G. haughtont Cooke 1949): In the description of the tooth
(Section II, chapter 2), statistical data was presented to support the hypothesis
that this specimen is not a lower molar. This may be strengthened further by
comparing the specimen with known upper Sivatherine premolars, for example
F 2989 and 4025 (fig. 25). It would not be difficult to reconstruct three roots
on the specimen; its base fits perfectly and directly on F 2989, which is worn
right down to the crown-root junction. The A—P axis and the breadth of F 39
are smaller than those of F 2989 and 4025. Comparison with other upper
524 ANNALS OF THE SOUTH AFRICAN MUSEUM
premolars indicates that from the points of view of other dimensions and of
general appearance, F 39 may be considered as an upper premolar, probably
P?. ‘The major objections to this diagnosis are:
(1) the extreme hypsodonty of the specimen, and
(2) the bulge on the lingual surface just below the crown-root junction.
The latter may be partly explained by the X-ray appearance (vide supra)
and partly by individual variability. In connection with the hypsodonty, it is
unfortunate that the available upper premolars of all the Sivatherine specimens
are so few, and also that they are in advanced stage of wear. Consequently an
adequate range of variation cannot be obtained for comparison. However, if
one attempts to reconstruct available specimens such as 4025 then it would
appear that the hypsodonty of F 39 may be equated. An unsatisfactory feature
of this diagnosis is that the upper molar Old. 109—an almost unworn M?—
measures only 47-0 mm. Even if this difference of 20 mm. (the teeth are at
approximately the same stage of wear) were decreased by a greater number of
specimens widening the ranges of variations, there would still be a significant
difference in height. However, it is important to consider the dentition as a
whole in the skull. If one examines the fairly complete S. giganteum skull
(15283) in B.M.N.H., it is seen that there is a distinct downward convexity in
a mesio-distal direction between P? and M3. Furthermore it is seen that P*
is approximately at the summit of the convexity and consequently it is expected
that P? and P* should be more hypsodont than M3. It is thus reasonable not to
exclude this specimen from S. olduvaiense only on the basis of its hypsodonty,
particularly because it is a single specimen and the comparable series is small.
However, it is felt that there are sufficient grounds to tentatively place the
specimen in a subspecies, namely haughtont.
Because Cooke (1949) compared this specimen with Sivathertum (Griqua-
therium) cingulatum, it is necessary to comment on his remarks. The bases of
distinction drawn by Cooke for separating F.39 from Sivathertum (Griquatherium)
cingulatum are not considered to be sufficient for a specific difference. First, the
central pit of a tooth is so variable that it cannot be used for differentiating two
species. Secondly, the small piece of cingulum remaining visible on the tooth
indicates that the cingulum zs marked. For these reasons, and because of the
fact that these two specimens come from the same area, although the exact
localities of their discovery are unknown, the probability that these two unusually
gross teeth belong to the same species is not insignificant. This independent
conclusion supports the view stated above that S. cingulatum could be considered,
on a subspecific level of S. olduvaiense. Further discoveries are required to resolve
this problem.
IV. MAKAPANSGAT
It has been stated above (111 a) that specimens M 553 B! (G. cingulatum)
and M 553 B have now been referred to Sivatherium olduvaiense. ‘This also holds
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 525
good for M 553 A and the other Sivatherine material described and discussed
above (Section 11, chapter 3).
V. ORANGIATHERIUM
The horn-cores mentioned by van Hoepen (1932) show all typical features
of those of S. olduvaiense, from North Africa, Olduvai and Hopefield. However,
it has been mentioned that it is distinctive only for its extremely grooved
appearance on its antero-medial convex surface. There are slight variations in
regard to the position of the knobs, but these are known to be highly variable
features.
The four teeth from the same site (C 426 A, B, C and D)—two of them
showing contact surfaces, and all four presenting the same degree of wear—
almost certainly belong to one individual. Lack of accurate information con-
cerning their discovery makes it impossible to associate definitely the teeth with
the horn-cores. However, it would seem likely that they belong to the same
species, if not the same individual, because they are derived from a single farm.
Furthermore, van Hoepen (1932) stated that the teeth most likely belong to the
horns. From a morphological point of view, C 426 A, B, C and D are identical
to those extreme stages of wear in the S. olduvaiense material, but there are some
important metrical differences. C 426 A is longer and broader than the M? of
either Hopefield or Olduvai. Although the length of the anterior and posterior
pillar do not differ much from S. olduvaiense, the great enlargement of the
metastyle in C 426 A provides it with a great increase in length. The metastyle
has been noted to be a well-developed character: but it is even more prominent
in this specimen than in MMK 3685.
Owing to the fragmentary nature of B, C and D, nothing can be said of
the styles, but the length of the pillars, as for C 426 A, fall beyond the dimen-
sions of the few M available.
Unfortunately, the teeth are in extreme stages of wear, even the bases of
the roots showing signs of attrition, and little more can be said of the distinctive
features of these teeth.
Because the horn-cores are so similar to those of S. olduvaiense (in which
some specimens have marked grooves) and because the few morphological
features of the dentition are also similar to S. olduvaiense, the authors hesitate
to provide a new species only on the basis of dental size, especially in the light
of the fact that so few M® are available and that the growth pattern in these
few teeth appears to show a marked tendency towards ‘abnormalities’.
It would appear that, because of the wide range of variation within the
species, gross dental metrical exaggerations should be considered on a sub-
specific level until clear criteria for specific differences can possibly be elicited
in specimens yet to be discovered. These dental maxima may only be ecological
variations, but, on the other hand, future evidence may indicate these maxima
to be of a specific nature.
526 ANNALS OF THE SOUTH AFRICAN MUSEUM
Consequently, even though criteria for a ‘subspecies’ in fossil material are
difficult to elicit and clarify, it is proposed to refer the specimens of Orangiatherium
vanrhyni to a subspecies of Sivatherium olduvaiense. ‘The question of priority which
would normally arise both as regards genus and species, falls away as van
Hoepen’s terminology is invalidated under Article 25(c) of the International
Code of Zoological Nomenclature. To link the specimens with discoverer, it is
proposed to designate them Sivatherium olduvaiense vanhoepent.
The plaster cast representative (C.1492) from Florisbad appears to fall
within the range of variation of Sivatherium olduvaiense.
VI. ?CORNELIA, ?FLORISBAD
The two milk molars marked B, and B, are Sivatherine in form and size
but nothing is recorded of the site of their discovery. Because of lack of com-
parative deciduous teeth and because of the fact that more than one subspecies
might be represented in the Orange Free State, the authors have decided to
refer these two teeth to an indeterminate subspecies of Szvathertum olduvaiense.
ConcLuDING NoTE on TAXONOMY
It is now necessary, on the basis of the above discussion, to modify Colbert’s
classification (1935) of the genera of the Sivatheriinae as follows:
Stvatherium syn. Indrathertum
Bramatherium
Hydaspitherium
Helladotherium
Vishnutherium
Libytherium
As far as the African fossil Sivatherine material is concerned, the following
genera, species and subspecies are recognized from 22 sites extending from
North Africa to the southern tip of South Africa:
Libytherium maurusium Pomel 1892
Stvathertum olduvaiense Hopwood, 1934.
Stvatherium olduvaiense haughtoni
Sivatherium olduvaiense vanhoepent, subsp. nov.
Stvatherium olduvaiense subsp. indet.
Stvathertum cingulatum
CHAPTER 3
THE FAUNAL RELATIONSHIPS AT THE AFRICAN
SIVATHERINE SITES
All the available data concerning the fauna identified at the various
African sites (figs. 17a, 18, 23) where Sivatherines are known to have been
527
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
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MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 531
discovered is summarized in table 60. No material from the Vaal River deposits
is included in this table because the Sivatherine specimens MMK 3685 and
F 39 were recovered from unknown localities, and it would be impossible to
associate them with any particular faunal group from the Vaal River as the
fauna are derived from series of gravels of varying geological periods, the
sequences of which have not been finalized. The data concerning the O.F.S.
sites is incomplete, because in the case of Tierfontein very little is known of the
associated fauna, while the mass of material collected from Florisbad and
Cornelia has not been identified.
It will be noted that only the genera are given in the table, because in very
many cases there is dubiety about the specific identification. Furthermore, in
some cases there is controversy concerning the genera themselves and some have
been referred to or included in others (Hopwood and Hollyfield, 1954).
Arambourg (1947) and Leakey (1958) differ, for example, as to the generic
determination of the East African suidae. Purely on a statistical basis, the
genera common to two or more sites have been selected and expressed in a table
of correlation (table 61).
( ~
ee al ree: emits ep
Se ae ee i ye ee
eee MOmiee aye, oe
Rh SS iy ion eS) Ge g 4 a 3s Oe Oe, te
Se CunemnOn © HOMO May. Miweh ik Oo
Garet Ichkeul 9 4 6 Gi 4 7 3 6 4 4 5 Sa ope T
St. Arnaud 2 ene 7 mea 40) 6) 11 5 7 7 5 5) Eye ish 1 te
Wadi Natrun 3 2 2 2 2 3 2 2 3 Teer otras I
Olduvai I 2Oie) Ui LF OR eure) | TORN ary: On 2a
Olduvai II Lea 2 2y hy RO Mh Say HP Say GLA ee tO” |) 4c 4
Olduvai ITI 12 8 Gi 6 6 8 Ria eal o
Olduvai IV Ieee Ay eS Fe 2Oun EAN 5. =e A
Olorgesailie 10 8 6 8 Sy gSte a Ee}
Omo II a | TA ORAL Ges
Serengeti 8 12 SOAP ato
Kanam 6 (i GIR)
Makapansgat OES ee ow
Hopefield 6 2
Florisbad 3
‘TABLE 61. Genera common to two different African sites.
(See also Appendix.) Since this table was drawn up a number of extinct genera have been
identified at Hopefield and at Olduvai, but they have not been included in this table.
The interpretation of such a table must be undertaken with great care
because of the varying amount of material collected from different sites. This
table does not express a relative time correlation. However, if one takes
Olduvai, Omo, Makapansgat and Hopefield, where large numbers of specimens
have been recovered and identified, the correlation becomes more representative,
but the actual figures do not indicate which genera are in common to all sites.
A similarity between two sites is indicated: Hopefield for example appears to
532 ANNALS OF THE SOUTH AFRICAN MUSEUM
have more in common with Olduvai IV than with Omo. Furthermore, if one
estimates the extinct forms and the extant forms at each site, and works out the
index of the relationship between extinct forms and the total, and between
extinct and extant forms, one obtains an interesting gradation of series;
especially if one selects arbitrarily those sites where there are more than a total
of 15 recognized genera (table 62). For both indices one obtains the same
gradation of increasing indices, namely, Hopefield, Olduvai IV, Serengeti,
St. Arnaud, Makapansgat, Olduvai II and Olduvai I. It must be emphasized
that this is not an attempt at an age sequence, but merely indicates the propor-
tions of the extinct and extant fauna. Nevertheless there is some evidence that
Hopefield overlaps the Olduvai IV period, and some of the faunal evolutionary
sequences (e.g. higher crown of Mesochoerus lategani (Singer & Keen, 1955)
compared with Mesochoerus olduvaiensis) and the more evolved human-
manufactured stone tools (Singer & Crawford, 1958a), tend to corroborate this
and even suggest that a portion of the ‘Hopefield period’ extends to slightly
more recent times than the period indicated by Olduvai IV.
Number of recognized genera
: Extinct Extinct
Total Extinct xX 100
Total Extant
Garet Ichkeul 13 3 Dat 30:0
St. Arnaud 16 4 25°0 BE)
Wadi Natrun 9 4. 44°5 80-0
Olduvai I 28 IO 35°7 55°6
Olduvai II 28 9 32-0 47°5
Olduvai III 13 5 38°4. 62°3
Olduvai IV 35 7 20:0 25°0
Olorgesailie II 4 36-4 57°0
Omo 26 9 34°6 53°0
Serengeti 33 8 24°3 32°1
Kanam 14 6 42°9 75°0
Makapansgat 48 15 Big 45°5
Hopefield 30 5 16°7 20°0
Florisbad 14 I 7% 749
Cornelia 7 4. 57°2 132°6
TABLE 62
However because all the material from Olduvai, Makapansgat and Hope-
field has not yet been definitely identified, these indices may have to be
altered in time (see ‘Appendix’). The data indicate clearly how, throughout
Africa, a large percentage of the fossil material is extant and how these extant
forms are widespread throughout the African Pleistocene. Consequently, it is
not surprising that Giraffa has been recovered from Upper, Middle and Lower
Pleistocene sites, and that Sivatherium has been recognized in each of the four
Beds at Olduvai and even at Omo.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 533
Confined to
Confined to Confined to
Stage I Stage IT Stage III
— Common to —————— Commonto =——————
Omo- I-II Olduvai I-II II-III Olduvai III-IV
Kanam Serengeti Olorgesailie
Dinopithecus Crocuta FAystrix Stmopithecus Otocyon
Lepus Anancus Aonyx Canis Bubalus
Syncerus Deinotherium Acinonyx Mesochoerus Nesotragus
Aepyceros Sus Chalicotherium Bularchus Philantomba
Kobus Metridiochoerus Pultiphagonides Adenota Phenacotragus
Antidorcas Mammuthus Parmularius Hippotragus Redunca
Menelickia Serengeticeros Gorgon Pelorovis
Omochoerus Serengetilagus Beatragus Damaliscus
Stegodon Heterocephalus Felis Thaleroceras
Archidiskodon AXerus Notochoerus
Homotherium Pedetes Choeropithecus
Tachyoryctes Equus
Mungos
Orycteropus
Metaschizotherium
HAylochoerus
Okapia
16°:2% 8-8% 250% 176% 13°2%
Common to Stages I, II and III
Hyaena Phacochoerus
Hipparion Hippopotamus
Ceratotherium Giraffa
Diceros Alcelaphus
Potamochoerus Oryx
Gazella Strepsiceros
T aurotragus
1G. 5
Total: 68 genera.
TABLE 63. Correlation of East African Pleistocene Fauna.
A correlation of the East African fauna as it is presently described, according
to the stages recognized (table 63), produces a number of genera (20°) common
to all three stages, which is almost identical to incidences in the Hopefield
material. ‘This indicates that throughout Africa about 20 per cent of approxi-
mately 70 genera persisted (though possibly undergoing specific determination)
over a great length of time, despite changing ecological and climatological
conditions. Furthermore, in respect of a single genus, namely, Giraffa, and one
single species, camelopardalis, which has been recovered from the Lower Pleisto-
cene (at Omo, see Arambourg 1947), there has been no evolutionary change,
despite its wide dispersal—chronological, climatological and spatial. This fact
demonstrates the tremendous adaptability of G. camelopardalis. However, during
this period, a much smaller species, G. gracilis, became extinct, as well as two
genera (Sivatherium and Libytherium) of another subfamily (Sivatheriinae).
534 ANNALS OF THE SOUTH AFRICAN MUSEUM
APPENDIX
After the MS. had been completed, three series of giraffid material were
made available to the authors. Newly discovered specimens from the Lime-
works breccia at Makapansgat were kindly sent by Mr. J. W. Kitching from
the Bernard Price Institute for Palaeontological Research, Johannesburg.
The Curator of Vertebrate Palaeontology (Dr. A. J. Sutcliffe) of the British
Museum (Natural History) informed us that some more East African fossil
giraffid specimens had been found in storage which had previously not been
known to be available. The third series, one specimen, was discovered by one
of the authors (R.S.) at Baard’s Quarry, Langebaan (Cape Province), which
is about 10 miles NW. of the Elandsfontein site at Hopefield. This specimen, an
astragalus, had been recovered, with other bones, from the layer of phosphatic
nodules about 5-10 feet below the surface. Most of the bones have been identi-
fied and they belong to animals similar to those found at Elandsfontein. Only
one identifiable specimen, Stegolophodon sp., belongs to a much earlier horizon
(Singer & Hooijer, 1958). |
A. MAKAPANSGAT
MATERIAL
I. Giraffa
M 2085: Left M, or M,.
M 1801: Right canine with tip of root broken off.
M 1798: Portion of left ramus of mandible with M, and fragmented Mg.
M 1800: Fragment of maxilla with portions of right M}!, P4.
IT. Sivatheriinae
M 2087: Fragment of mandible containing unerupted I,, I,.
M 2086: Fragmented right M3.
M 539 A: Now in British Museum (Natural History) and numbered
M 16729. Jaw fragment of juvenile.
DESCRIPTION
I. Giraffa
M 2085
This is a left lower molar, probably M,, but the possibility of its being M,
cannot be ruled out. It is a complete tooth with a fragment of mandible between
its roots. The crown is identical in appearance to M 942 —M 1113, except that
(i) M 942 has a minute ectostylid and M 2085 has no trace of it;
(ii) the stylids on the lingual surface of M 2085 are less marked than those
of M 942—M 1113; 7
(iii) M 2085 is in a more advanced stage of wear; and
10
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 535
(iv) the posterior pillar of M 2085 is more rounded at the base of the buccal
surface than M 942—M 1113, and M 2085 does not present the same
marked indentation of the crown on the posterior surface that M 942 —
M 1113 has.
The roots of M 2085 are very robust, being more massive than those of
the modern G. camelopardalis, although they are as short.
Height
CROWN Length Breadth Lingual Buccal
(a) Anterior pillar 15°5 25°5 19°6 16°3
Posterior pillar E77 25°0 18°5 14°3
Whole tooth 34°7 25°5
(d) Roots Breadth Height
Anterior 24°4 28°8
Posterior 23°7 28-0
Table 64. Measurements of M 2085 (mm.).
These dimensions fall within the range of the other Makapansgat fossil
Giraffa; the breadth falls within the range of variation (at upper end) of the
modern Giraffa camelopardalis while the length falls just outside the range of the
modern species.
This, like the other Makapansgat Guraffa specimens, is included in
G. camelopardalis.
M 1801 (PI. 23, e, f)
This is a rather worn right canine with most of the root intact. It presents
no features not present in a canine of a modern G. camelopardalis. Its dimensions
fall within the range of the modern species:
Crown: Length: 21-5 (mm.)
Breadth: 10-2
Height: 18-0
Root: Base-tip: 33+
M 1798
A portion of the left horizontal ramus of a mandible with a complete M,
and a portion of the anterior pillar and the entoconid of M,. The buccal
portion of the mandible is partly broken away exposing the anterior root of M,
and the sockets of the roots of P,. The teeth are in a fairly advanced stage of
wear, intermediate between the stages of M 942—M 1113 and M 2085. The
general appearance of M 17098 is similar to that of the other Makapansgat
fossil Giraffa M, specimens, except that it has a prominent ectostylid and only
a slight hypostylid which has been worn away by the abutting anterior pillar
of M,. The anterior root of M, tapers towards the tip, in contrast to that of
M 2085 which is rectangular.
536 ANNALS OF THE SOUTH AFRICAN MUSEUM
This specimen is the only M, of the fossil G. camelopardalis in the present
survey.
The remaining portion of the M, of M1798 resembles the other M,
Makapansgat specimens, being tightly wedged against the posterior pillar of
M,. On the lingual surface of the entostylid the median costa is dimpled by a
V-shaped vertical depression.
(2) ManprsLe: Breadth opposite M,/M,: c. 34
Height opposite P,/M,: ¢. 54
Height
M, Length Breadth Lingual Buccal
(6) Anterior pillar 15°5 22°79 c. 14 14°9
Posterior pillar 16°3 23°4 c. 15 14°2
Whole tooth 31°6 22°77
M,
Anterior pillar 16°3 G29 c. 20 18-0
Posterior pillar _ — 20°6 —
TABLE 65. Dimensions of M 1798 (mm.).
M 1800
A fragment of right maxilla containing portions of P* and M!. They are
in a very advanced stage of wear, the most worn down of the Makapansgat
Giraffa series.
M, Length Breadth Height
Lingual Buccal
Anterior, pillar. 3411-5 29:0 7:0 6:5
Posterior pillar... — — - 8-2
Whole tooth .. 25+ — —- —
Dimensions of M 1800 (mm.)
IT. Stvatheriinae
M 2807
A fragment of the right body of a Sivatherine mandible, the symphyseal
aspect of which fits that of M 553A perfectly (Pl. 40). A small portion of the
posterior border of the body is present forming an arch with that of M 553A,
but an anterior directed fracture has separated the body from the horizontal
ramus about I cm. along the posterior border to the right of the symphysis.
The anterior half of the mental foramen is present, as well as a small accessory
foramen 11 mm. beyond the anterior border of the mental foramen with which
it is continuous by a canal in the bone. Anteriorly the unerupted crowns of
I, and I, are visible where the outer bony alveolus has been broken away and
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 537
the socket for the root-tip of I, is visible. It is evident that I, is at a more
advanced stage of the process of eruption than I, which is ‘impacted’ against
I, and is partly overlapping it. The anterior enamel edge of the crown of each
tooth is at an angle of about 45° to the superior surface of the body of the
mandibie, the medial edge of each enamel border being nearer the surface.
Furthermore it is obvious that I, considerably overlapped the plane of I,,
indicating that during subsequent growth considerable lateral expansion of the
body was still to occur so as to accommodate all the teeth (see also p.498).
It is curious that there is no sign of the canine, although the broken edge of the
mandible just lateral to I, appears to be the socket for the root. On the left
this region is obscured by breccia. The order of eruption, as regards the
premolar-incisor-canine teeth, then, simulates that in modern Giraffa camelo-
pardalis. ‘The enamel of the incisor is fairly rugose and the crown enamel is
markedly convex, the occlusal edge of the enamel projecting vertically. If one
mentally reconstructs the anterior portion of the body one obtains the conviction
that the incisor would not be in the same plane as the superior surface of the
body but rather at about 45° toit. This tends to confirm our opinion stated on
page 408 that the incisor-canine row of the Makapansgat Sivatherium olduvaiense
would be at an intermediate position between Hydaspitherium sp. (AMNH
19684) and Giraffa camelopardalis.
It is now possible to assess more accurately the length of the body of the
mandible, namely, about 120 mm. This specimen also indicates that the height
of the body of the mandible relative to the breadth of the body is greater in the
immature individual. With maturation of the individual the depth decreases
and the breadth increases. This principle also pertains to the modern G.
camelopardalis.
The only measurement that can be taken on the teeth is the breadth of the
crown of I,, namely, 23-7 mm.
M 2086
This is a right M;, broken off at the crown-root junction with the anterior
pillar almost complete, the posterior pillar having the occlusal portion of the
hypoconid broken off, and the talonid having most of the buccal cone broken
away. The slightly rolled edge of the enamel of the occlusal surface indicates
that the tip of the crown of the tooth was only just above the alveolar margin of
the mandible, so that most of this specimen must still have been embedded in
the mandible. This very early stage of eruption is in conformity with that at
which a M, of M 553B* would have been expected to be. Furthermore, as was
found with M 2087 which belongs to M 553A, it would be reasonable to con-
clude that M 2086 is the M; of the right side of the individual to which M 553B},
M 553B, M 553 and M 2087 belong. It can also be concluded that this specimen
supports the view that the order of eruption of the molars of Sivatherium is the
same as that in Giraffa camelopardalis, namely, M,-M,—M, in that order.
538 ANNALS OF THE SOUTH AFRICAN MUSEUM
It is interesting to note that in M 2086 the angularity of the buccal cones
seen in M 553B? is obvious only in the second pillar of M 2086 while its anterior
pillar is more rounded. |
In general appearance, M 2086 closely resembles M, of Old. SK. II, 92,
except that the parastylid of M 2086 is a more marked ridge, while the median
ridge (costa) of each pillar of M 2086 is not yet as prominently developed as
those of the Olduvai specimen (which is at a much later stage of eruption).
The finely rugose buccal surface of M 2086 has three horizontal ridges on the
enamel on the anterior pillar and one on the posterior pillar—these are more
marked than in Olduvai 92.
Measurements of M 2086 (mm.) :
Length Breadth Height
Lingual Buccal
Anterior pillars. |) 26-0 33:0 49+ 49+
Posterior pillar .. 22-0 29+ 5r+ —
Talonid .. .. 14+ 19+ 31+ —
Whole tooth .. 66+ 33-0 — =
It is clear that this specimen falls within the range of the measurements of
Sivatherium olduvaiense, and, being the least worn of the Msg series, it extends the
upper range of variation of the height of the tooth to 51-++ mm.
M 539A (PI. 36, g)
This is the number given by the Bernard Price Institute for Palaeonto-
logical Research, Johannesburg, but it isnow permanently in the British Museum
(Natural History) where it has been given the number M 16729.
It is a fragment of a left mandibular ramus of a juvenile and definitely
belongs to the opposite side of the Sivatherine specimen M 539B. It contains
the two posterior pillars of DM4, the most anterior pillar being broken
away.
Measurements of mandibular fragment (mm.) :
Menetiiat een aoe:
inickitessi ace ies
B. ADDITIONAL East AFRICAN SPECIMENS
The additional giraffid specimens in the British Museum (Natural History)
are mainly fragmentary and are derived from Olduvai Gorge (Tanganyika),
Laetolil beds (Vogel River, Tanganyika) and Broken Hill (Northern Rhodesia)
(figs. 17a, 17b). Brief descriptions of the Sivatherine specimens only are
given.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA
(a) OLpuvart GorGE
MATERIAL
I. Giraffa
M 14778:
M 14781:
M 14702:
M 14793:
M 14794:
M 14795:
M 14706:
M 14797:
M 14708:
IT. Sivatheriinae
M 14535:
M 14779:
M 14780:
M 14701:
Axis. Bed I.
Cervical vertebra (7th). Bed I.
Calcaneum. Bed I surface.
Metacarpal, distal end. Bed I.
Metatarsal, distal end. Bed I.
Metacarpal, distal end. Bed IT.
Metatarsal, distal end. Bed II.
Metatarsal, distal end. Bed II, surface.
Metacarpal, proximal end. Bed I.
Horn core fragment. Bed I.
Cervical vertebra (6th). Bed IV.
Cervical vertebra (6th). Bed I, surface.
Tibia. Bed ITI.
M 17024-6: Three molar teeth.
?: Unnumbered left lower molar. G. RK III.
III. Giraffid (no generic distinction possible)
No numbers on specimens. Letters refer to sites.
Astragalus .. Oldy. FLK II 8
— 10] Cited!
~ HEK II 8S
a HEK II S
— GTS IVS
—~ GRK II S
Phalanx! ). )...:) Oldy. GHJK IIS
— EHK I
_ DK. 15s
_ HEK II
Calcaneum Dy 1 GRK eS
Tibia Oldy. FC II S_ (distal end)
— EK I S_ (proximal end)
_ Sc II S_ (proximal end)
— GRK II S_ (proximal end)
— DK. ;.1 base
Cubonaviculare
(Scaphocubeid)), Oldy.. THC 1, (y)
— VEK I
= SHK II 8
Sas)
oa°
Metapodial
ANNALS OF THE SOUTH AFRICAN MUSEUM
Oldy. FC II S_ (distal end)
— DET Ss tdistalend}
~ HEK I (distal end)
(b) Lartoxit BEps
I. Stvatheriinae
M 15088: Lower molar (or premolar).
M 15089: Upper premolar.
M 15090a, b: Lower premolars.
IT. Giraffid
Cubonaviculare
(Scaphocuboid) 1710.5
Lit.A.S.
Phalanx 1’ MnxX.S.
LIT.AS.
Astragalus .. 17108
LIT.A.
(c) BRokEN Hii
I. Giraffa
Ni 2126,
ts
Il. Sivatherwinae
M 12128 ..
M 12129 ..
III. Giraffid
Tibia, distal end.
Unnumbered horn-core, cervical vertebra and distal end of
a femur.
Metacarpal, distal end. (Same number as Giraffa tibia.)
Astragalus.
Radius, distal end.
DESCRIPTION OF SIVATHERINE SPECIMENS
M 14535
This fragment of horn core is entered in the record book as being derived
from ‘Middle Pleistocene Bed I Olduvai, Tanganyika, Leakey collection
1932".
One side is slightly convex in an A’-P direction and has 3 deep, wide grooves
(7-8 mm. in breadth) more or less parallel to each other and to the length axis
of the horn. The other surface is concave and relatively smooth.
Although it is a small fragment it is very similar to the South African
specimens, It is typically Sivatherine.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 541
M 14791
This is a complete tibia. Its dimensions are compared with the average of
5 Giraffa camelopardalis from Central Africa in the Musée Royal du Congo Belge
in Tervuren.
Giraffa
M 14791 camelopardalis
Total length Ks : 473 639
Proximal end: Transverse 153°5 151°8
A-P bie 138-0 Q2°4
Mid-shaft: Transverse 64°3 70°6
1 a Rds 49°4 55°8
Distal end: ‘Transverse IO1°O 109:0
A-P ee 75°4 76:2
TABLE 66. Measurements (mm.) of M 14791
compared with Giraffa camelopardalis.
This again bears out the contention expressed above that Sivatherium
olduvaiense is much shorter and has more massive extremities than the modern
giraffe.
M 17024-26
These 3 teeth of Sivatherium olduvaiense are all derived from Bed II.
M 17024
This is a right lower molar, either M, or M,. The posterior end of the
posterior pillar has been broken away. It is in early wear and the enamel is
very rugose. It is markedly hypsodont and the cingulum bulges slightly. The
costae are very prominent, as are also the protostylid and metastylid. The roots
are broken off.
Measurements (mm.) of M 17024:
Maximum breadth .. ise SA ey PS
Occlusal breadth, anterior pillar .. 17:8
posterior pillar .. 17-9
Height, lingual ye i) ta AD
buccal a Me Se) ne Ord:
M 17025
This right upper molar is probably M?. The anterior pillar is partly
broken away on the buccal side. The enamel is coarsely rugose. The lingual
surface slopes markedly from the base in a buccal direction. The styles are
very prominent. The roots are broken off.
Measurements (mm.) of M 17025:
Maximum length .. 49
Maximum breadth .. 43
Occlusal length SH Wheel oho
542 ANNALS OF THE SOUTH AFRICAN MUSEUM
M 17026
This is a left M! or M? in advanced wear. There is no cingulum. There is
a marked protostyle and the median costa of the anterior pillar is flattened.
Most of the posterior pillar is missing and the 2 buccal roots are broken off.
The lingual root forms a broad plate.
M 15088-15090
These Sivatherine teeth are from the Laetolil Beds (Vogel River) and
recorded as ‘Pleistocene Tanganyika. Leakey collection 1935’. M 15088 and
15090a, b are lower P, and M 15089 is an upper premolar.
M 15089
This right upper premolar has its crown fairly well preserved. It is finely
rugose and. the lingual surface has a marked slope. Wee median costa is very
prominent. The root is broken off.
M 15090a
Its anterior root is broken away and belongs to the side opposite that of
15090b.
M 15090b
The anterior pillar is worn down and it projects much more than the
posterior pillar. The enamel is very finely rugose. The buccal surface of the
anterior pillar is missing and the roots are broken off.
Length (A—P) Breadth Height
Max. Occlusal Max. Occlusal Buccal Lingual
M 15088 BO 36-2 33°0 20°0 30°6 32°5
M 15089 33°2 29'°8 46-7 33°3 29°8 26-0
M 150904 37°8 3555 27°38 — 13-1 21°4
M 150906 36:3 36-2 32°0 28-7 24:6 30°1
TABLE 67. Measurements (mm.).
Unnumbered GRK II Olduvai
This is a lower left molar, probably M,. It is still embedded in matrix.
The stylids are very prominent.
Measurements (mm.) :
Maximum length .. 47
Maximum breadth .. 49
Occlusal breadth, 7") 38
Height si et a Monee
M 14778
This is an axis. The total height is 172 mm. and the total A—P length is
160 mm. The neural canal measures 40 mm. transversely and 38 mm. A—P.
The inferior articular surface measures 65 mm. transversely and 77 mm. A—P.
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 543
M 14779
This is a 6th cervical vertebra. The maximum breadth of the body is
132 mm., and the inferior articular surface measures 74 X57 mm.
M 14781
This is a 7th cervical vertebra from Bed I at Olduvai. The maximum
breadth of the body is 115 mm., the neural canal is 45 mm. in diameter and
the inferior articular surface is 104. mm. broad and 76 mm. A-P. The condyle
(on superior aspect of the body) measures 67 X 51 mm.
M 12128 and M 12129
These are specimens from Broken Hill, Northern Rhodesia. It is not certain
whether these are Sivatherine.
M 12128. This is a distal end of a metacarpal and measures 117 X 74. mm.
M 12129. This is an astragalus and measures 81 X 122 mm.
The following giraffid postcranial specimens are from the Olduvai Gorge.
They are not numbered and no generic distinction is possible. Only relevant
measurements (mm.) are given.
(i) Astragalus
Max. length A-P ‘Transverse
INovsite ).. oe 106 60 70
Oldy FLK IIS... 103 61 65
tl DOs i Weare IOI 54 63
» HEKOILS .. Veit 67 73
oe el EIS Te Sic, 102 63 65
PG Gi UN Sales IOI 62 63
5 GRE IPS... 106 62 70
(ii) Phalanx I :
Maximum Shaft Prox. end Distal end
length A-P Transv. A-P Transv. A-P Transv.
Oldy GHJK IIS .. — 104 44 54 B58 Bus 55
Pee lis * ie 121 — — 56 59 37 53
Pe DI TS. 114 AP 52 Som 59 Ba) 55
Peedi DEN 109 44 46 54 554 a7 50
(111) Calcaneum
GRK IIS Maximum length .. 222
Body) tansy) iach
height ue aes OO
oeee ANNALS OF THE SOUTH AFRICAN MUSEUM
(iv) Tzbza
Distal end fragment: Oldy FC IIS ‘Transv... 97
ASP hit 86g
Proximal end fragment (without fibular style) :
Transverse A-—P
EK. ALS oy 110 74
SC ELS 105 69
GRKIIS 87 77
DK I Base 87 81
(v) Cubonaviculare (scaphocuboid)
Thickness A-P Transverse
Oldy, THG, 1.W). 42 76 103
sie MKT Hie 43 77 IOI
3 be FS haar 36 64 64
(vi) Metapodial
Distal end fragments: A-P ‘Transverse
EQ Se 50 93
DIRS a Sne a Aiea 65 112
HEK I Me Me 57 105
The following giraffid postcranial specimens are from the Laetolil Beds
(Vogel River, Tanganyika):
(i) Astragalus
Max. length A-P ‘Transverse
LO ee 113 58 62
PAS AY 8G. 102 58 62
(ii) Phalanx I
Maximum Shaft Prox. end Distal end
length A-P Transv. A-P Transv. A-P Transv.
MnX S_.. 97 39 35 48 49 43 30
LIT.AS .. 104 bie, 445 47 49 39 27
(* denotes exostosis included)
(iii) Cubonaviculare (scaphocuboid)
Thickness A-P ‘Transverse
1710.5 a 4.7 89 107
LT. Ae: 40 82 105
C. LANGEBAAN (CAPE PROVINCE)
S.A.M. 11715 (Pl. 50, c, d) |
This is an almost complete right astragalus (talus) now housed in the
South African Museum, Cape Town. It is longer than Olduvai 102 and 107,
MODERN GIRAFFES AND THE FOSSIL GIRAFFIDS OF AFRICA 545
but its general appearance is identical to theirs. It belongs to Sivatherium
olduvaiense.
Maximum proximo-distal length .. erie ae =e
Maximum A-P length medially .. “ 71
Maximum A-—P length laterally .. eo 26/67
Maximum breadth proximally... és 84+
Maximum breadth distally . . as ee 74.
Maximum articular breadth proximally .. ial
Maximum articular breadth distally f. 74
ACKNOWLEDGEMENTS
A. FINANCIAL AID
One of us (E.L.B.) received a travel grant from the United States
Educational Foundation and the Fulbright Plan Organization, and a research
grant from the Belgian Fonds National de la Recherche Scientifique (Brussels).
The other author (R.S.) received grants from the South African Council for
Scientific and Industrial Research and the Dr. C. L. Herman Research Fund,
University of Cape Town. In addition, each of the authors received research
grants independently from the Wenner-Gren Foundation for Anthropological
Research, Inc., New York. We are exceedingly grateful to these organizations
for the generosity which enabled all the fossil and recent material to be studied
and collected. Furthermore, the Land Rover donated by the Wenner-Gren
Foundation to the Hopefield Research Project of the University of Cape Town
was extensively used for field work.
We wish to record our thanks to the Director (Dr. A. W. Crompton) and
Trustees of the South African Museum, Cape Town, who have not only accepted
our paper for publication in a special issue of the Annals, but have also subscribed
a portion to its cost of publication. The major publication costs have been met
by special grants: (a) by the South African Council for Scientific and Industrial
Research and by the Council of the University of Cape Town, and (0) by the
Belgian ‘Fondation Universitaire’ to the other author (E.L.B.). For this
assistance we are extremely grateful.
B. Loan oF MATERIAL
The following very kindly released material (including type specimens)
on loan so that we could study most of the material in Cape Town: Dr. L.S. B.
Leakey, Curator of the Coryndon Museum, Nairobi (to whom we are particu-
larly grateful for his co-operation in sending all the East African fossils) ;
Professor R. A. Dart, University of the Witwatersrand; Dr. A. S. Brink and
Mr. J. W. Kitching, Bernard Price Institute for Palaeontological Research,
University of the Witwatersrand; Dr. A. C. Hoffman, Director of the Nasionale
Museum, Bloemfontein; Mr. B. D. Malan, Director of the Archaeological
Survey of South Africa; Mr. J. H. Power, recently Director, and Dr. R. Bigalke,
present Director of the McGregor Memorial Museum, Kimberley; Professors
546 ANNALS OF THE SOUTH AFRICAN MUSEUM
C. Arambourg and J. P. Lehman of the Muséum National d’Histoire Naturelle,
Paris; Dr. E. Colbert of the American Museum of Natural History, New York;
and Dr. A. J. Sutcliffe of the British Museum (Natural History).
C. DEPARTMENTs VISITED, TECHNICAL Alp, PHOTOGRAPHS
The individuals mentioned in ‘B’ are also thanked for giving permission
to study material (mainly recent) in their Institutions. In this respect we are
also grateful to Professor A. S. Romer and Dr. B. Lawrence of the Comparative
Zoology Museum at Harvard University (Cambridge, Mass.); Drs. G. G.
Simpson, J. Anthony and D. Carter of the American Museum of Natural
History; Dr. A. W. Crompton, South African Museum, Cape Town; Dr. W. D.
Turnbul of the Natural History Museum, Chicago; Dr. R. Kellogg, U.S.
National Museum, Washington; Dr. E. Schouteden, Musée Royal du Congo
Belge, Tervuren.
We thank Mrs. R. H. Nichols of the American Museum of Natural History
for her assistance and kindness; also Mr. Kenneth Abrahams, a medical student
at the University of Cape Town, who, with other students, greatly assisted on
field collecting trips. Radiographs were kindly made by Drs. B. and H.
Hirschon, Cape Town.
We are extremely grateful to Mr. G. McManus, Surgery Department,
University of Cape Town, who patiently made most of the magnificent photo-
graphs of the specimens, and to Miss L. A. Abrahams who typed the manuscript.
Mr. J. N. Darroch, Department of Mathematics, University of Cape Town,
gave helpful advice with some of the statistical analyses.
Miss A. Schweizer, South African Museum, kindly assisted with the
preparation of the plates.
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in 1957.)
Ann. S. Afr. Mus., Vol. XLV Plate I
a—Old. 2.53, antero-medial aspect. b—Old. 86, antero-medial aspect. c—Old. 1. 53, antero-
medial aspect. d, e, f—Old. 2.53, 1.53, 86 respectively, illustrating the described features on
their postero-lateral aspects. No scale.
Ann. S. Afr. Mus., Vol. XLV Plate II
Old. 3.53: a, c—postero-lateral and anterior views, respectively, illustrating features described.
No scale. b—antero-medial aspect.
Ann. S. Afr. Mus., Vol. XLV Plate III
a, b—postero-lateral and antero-medial aspects of M 14955 (Olduvai), respectively. c— Old. 1952
SHK II BK II base (S) plus M 149546, antero-medial aspect. d—idem, postero-lateral aspect.
No scale.
Ann. S. Afr. Mus., Vol. XLV Plate IV
scale em
1 2
Sonennssoncenannaciitennansccneenn tn tis erccnt TPO
scale em
1 a Os
erennnnnnninntnannnnanrcserennarnnnantennnnatt
F 3655 (Olduvai): a—buccal aspect. b—lingual aspect. c—occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate V
a— Old. 93, buccal aspect. 5—Old. F 3656, buccal aspect. c—Old. 93, lingual aspect.
d—Old. F 3656, lingual aspect.
Ann. S. Afr. Mus., Vol. XLV Plate VI
scale ver
Bi
a—Old. F 3656, occlusal aspect. b—Old. 93, occlusal aspect. c—Old. 1, buccal aspect.
d—Old. 1, lingual aspect. e—Old. 1, occlusal aspect.
Plate VII
Ann. S. Afr. Mus., Vol. XLV
scale om
} 2
a
Old. 6: a—lingual aspect. b—buccal aspect. c—occlusal aspect.
Plate VUI
Ann. S. Afr. Mus., Vol. XLV
a—buccal aspect. b—occlusal aspect.
Old. 365
Ann. S. Afr. Mus., Vol. XLV Plate IX
seale cm
1
Old. 392. a—buccal aspect. 5—lingual aspect. c—occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate
scale cm
1 ae
anannnconsccscssespaciods
Old. 3. a—buccal aspect. b—lingual aspect. c—occlusal aspect.
Ann. S. Afr. Mus., Vol XLV Plate XI
Old. 92: a—buccal aspect. b—lingual aspect. c—occlusal aspect.
*yoodse yesnjo00 —9
‘yoodse tensutt—q *joodse yeoonq—v :126 ‘plo
TIX #°1d ATX TA “SHY “APY *S “UU
*yoodse pesnjo00 — 9
‘yoodse jensulj—q ‘joodse peoonq—p ‘ows 1WUNSNy
TTX #4°1d ATX TSA “SAIL “FV “S “UUYy
Ann. S. Afr. Mus., Vol. XLV Plate XIV
a— Old. F 2993, buccal aspect. b— Old. F 2993, lingual aspect. c—Old. F 2993, occlusal aspect.
d—QOld. 109, buccal aspect. e—Old. 109, lingual aspect. f—Old. 109, occlusal aspect.
g—Marsabit Road, buccal aspect. h—Marsabit Road, lingual aspect. i—Marsabit Road,
occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate
AV
a—Old. 2, buccal aspect. b—Old. 4, buccal aspect. c—Old. 116, buccal aspect.
d—Old. 5, buccal aspect. e—Old. F 2989, buccal aspect.
Ann. S. Afr. Mus., Vol. XLV
Plate XVI
FS
BS
scale em
As in plate 15; lingual aspect.
Ann. S. Afr. Mus., vol. XLV Plate XVII
As in plates 15, 16; occlusal aspect.
Plate XVII
Ann. S. Afr. Mus., Vol. XLV
. 105, buccal aspect.
. surface’, buccal aspect.
. b—Old. 95, buccal aspect. c—Old
. e—Old, 120, buccal aspect. f—‘Old
a—Old. F 2991, buccal aspect
d—Old. 166, buccal aspect
Ann. S. Afr. Mus., Vol. XLV Plate XIX
scale em
a—Old. F 2991, lingual aspect. b—Old. 166, lingual aspect. c—Old. 105, lingual aspect.
d—Old. 95, lingual aspect. e—Old. 120, lingual aspect. f—‘Old. surface’, lingual aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XX
GY
yy
scale cm —
1 2 3
hoertesmerscommmmnastinrimemeascencsemeomnnmtoneritsusecsmmmmed,
As in plate 19: occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XXI
a—Old. 107, astragalus, anterior aspect. b—Old. 115, radius, articular surface (<— indicates
posterior border). c—Old. 101, tibia, anterior aspect. d—Old. 104, cubonaviculare, proximal
articular surface. e—
Old. 341, os magnum, distal articular surface. {— Old. 342, sesamoid bone,
side view. g—Old. 105, cuneiform, proximal articular surface.
Ann. S. Afr. Mus., Vol. XLV Plate XXII
a—Old. 114, metacarpal, anterior (dorsal) surface. )—Old. F 364, proximal phalanx, posterior
surface. c—Old. 185, proximal phalanx, side view. d—‘Old. surface’, proximal phal_nx,
side view.
Ann. S. Afr. Mus., Vol. XLV Plate XXIII
Bale Co
4 z
ceanerennneatnncnnnencteccomermecmteh
seale enn
a—Old. 103, calcaneum, medial aspect. b—Old. 103, calcaneum, lateral aspect. c— Old. 100,
femur, distal articular surface. d—Old. 116p, ulna, anterior aspect of proximal end.
e, {—Makapansgat M 1801; buccal, lingual aspects respectively.
Ann. S. Afr. Mus.,"Vol. XLV Plate XXIV =
a—Old. 106, metatarsal, posterior aspect of proximal end. b—Old. 106, metatarsal, anterior
aspect of proximal end. c—Old. 103, calcaneum, anterior aspect. d—Old. 314, metatarsal,
posterior aspect of distal end. e—Old. 314, metatarsal, anterior aspect of distal end. f—Old. 106,
metatarsal, proximal articular surface. g—Old. 104, cubonaviculare, distal articular surface.
Ann. S. Afr. Mus., Vol. XLV Plate XXV
MMK 3685 (O.F.S.): a, b—buccal and occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XXVI
O.F.S.), lingual aspect.
O.F.S.), lingual aspect.
F.S B
.F.S.), lingual aspect. d—C. 426C
4
c—C. 426D (
Ann. S. Afr. Mus., Vol. XLV Plate XXVII
a—C. 426A, buccal aspect. b—C. 426B, buccal aspect. c—F. 39 (O.F.S.), X-ray (antero-
-)
posterior). d—F. 39 (O.F.S.), posterior aspect. e—F. 39 (O.F.S.), buccal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XXVIII
seale em
‘1 2 6
a—C. 426A, occlusal aspect. b)—C. 426B, occlusal aspect. c—C. 426C, occlusal aspect.
d—C. 426D, occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XXIX
Gy
Z
a—C. 431B, lateral aspect. Note cranial sinuses in base. b—C. 431B, medial aspect.
c—C. 431A, anterior view. d—C. 431A, antero-medial aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XXX
scale em
1 2 3
‘eeecteeenmmnonieiteteenonsenannneenenneetennnnnndonneemmemeceneemeteell
scale em
1 2 4
‘remceceneeeennensinsnaenecsost enennetnarensereorsasuentnenenecieonearenenmeeetcoesennanaet
(O.F.S.), buccal, lingual, occlusal aspects respectively.
b, d, f—B? (O.F.S.), buccal, lingual, occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XXXI
Makapansgat specimens: a, i, g—M 646, buccal, lingual, occlusal aspects respectively.
b, j, r—-M 944, buccal, lingual, occlusal aspects respectively. c, k, s—M 536, buccal, lingual,
occlusal aspects respectively. d, 1, t—M 533, buccal, lingual, occlusal aspects respectively.
e, m, u-M_535, buccal, lingual, occlusal aspects respectively. jf, n, v—M 1115, buccal, lingual,
occlusal aspects respectively. g, 0, w—M 552, buccal, lingual, occlusal aspects respectively.
h, p, x -M 551, buccal, lingual, occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XXXII
seale em
1 2 &
J rvreerorrssnannnseslatsseneonomeneisensiencenr reso
Makapansgat specimens: a, b, c—M 525, lingual, buccal, occlusal aspects respectively.
d—M 938, buccal aspect. e—M 936, buccal aspect. f—M 1113 plus M 942, buccal aspect.
g—M 263, buccal aspect. h—M 528, buccal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XXXII
scale em
Makapansgat specimens, lingual aspect. a—M 938. b—M 936. c—M 1113 plus M 942.
d—M 263. e—M 528. f—-M 532. g—M 1114. h-M 531. i—M 264.
Ann. S. Afr. Mus., Vol. XLV Plate XXXIV
Makapansgat specimens, occlusal aspect: a—M 938. b—M 1118 plus M 942. c—M 936.
d—M 263. e—M 528. f—M 550.
Ann. S. Afr. Mus., Vol. XLV Plate XXXV
scale em
1 2 5
Makapansgat specimens: a, e—M 532, buccal and occlusal aspects respectively. b, f—M 264,
buccal and occlusal aspects respectively. c, g—M 1114, buccal and occlusal aspects respectively.
d, h—M_ 531, buccal and occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XXXVI
g
scale in cm
= OE Ds Be
Makapansgat specimens: a, b, c—M 527, buccal, lingual, occlusal aspects respectively.
d, e, f—M 1116, buccal, lingual, occlusal aspects respectively. g—cast of M 539A, now in
B.M.N.H. and numbered M 16729. Bucco-occlusal view.
Ann. S. Afr. Mus., Vol. XLV Plate XX XVII
scale em
4 Zz
g
YG
UG
Vy
Makapansgat specimens: a, h, k—M 939, buccal, lingual, occlusal aspects respectively.
b, g, i—-M 540, buccal, lingual, occlusal aspects respectively. c, e, j—M 937, buccal, lingual,
occlusal aspects respectively. d, f, /—M 524, buccal, lingual, occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XXXVIII
“4
scale in cm.
SS ee ee Se
Makapansgat M539B: a—bucco-occlusal aspect. 5—lingual aspect. c— X-ray.
Ann. S. Afr. Mus., Vol. XLV Plate XXX1IX
seale em
1 a 4
Makapansgat specimens: a, b—M 553A, buccal (lateral), lingual (medial) aspects respectively.
c, d—M 5538, lingual, occlusal aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate XL
scale em
1 os 4s
553A plus M 2087 (joined): a—shows buccal aspect of M 2087 and lingual aspect of
553A (inferior border adjacent to ‘a’). 5—inferior aspect. c—superior aspect.
419, J Pp Pp Pp
Ann. S. Afr. Mus., Vol. XLV Plate XLI
scale em
Leccnsretminininrcencnttarsisastorancnoeiptesatstet Meee
M 553B!: a—bucco-occlusal aspect. b—buccal aspect.
| Ann. S. Afr. Mus., Vol. XLV Plate XLII
<<
Hopefield specimens: a—4372B, viewed from endocranial aspect. b—4372B, inferior aspect:
shows incomplete occipital condyles. c—4373A, side view. d—4373A, opposite side to (c).
Ann. S. Afr. Mus., Vol. XLV Plate XLUI
Hopefield 4372. a—postero-lateral aspect. b—anterior aspect. c—antero-medial aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XLIV
Hopefield 4373: a—antero-medial aspect. b—anterior aspect. c—postero-lateral aspect.
Plate XLV
Ann. S. Afr. Mus., Vol. XLV
Hopefield specimens: a, f, g—4025, buccal, lingual, occlusal aspects respectively. b, e, i—4026,
buccal, lingual, occlusal aspects respectively. c, d, hR—3345, buccal, lingual, occlusal aspects
respectively.
Ann. 8S. Afr. Mus., Vol. XLV Plate XLVI
Hopefield specimens: a— 4027, buccal aspect. b—4023, buccal aspect. c—4024, buccal aspect.
d—4374, buccal aspect. e—4028 plus 4028A (considered as 4028), buccal aspect.
Plate XLVII
Ann. S. Afr. Mus., Vol. XLV
1 aspect.
ingua
b)
As in plate 46
Ann. S. Afr. Mus., Vol. XLV Plate XLVUI
SS
\
As in plates 46, 47; occlusal aspect.
Ann. S. Afr. Mus., Vol. XLV Plate XLIX
scale em
A 2
3
bneesemeeeestnncetmineeesenteenanannensbeereentmnmanantenad,
scale em
bette etree nena never!
Hopefield specimens: a, b, c—4029 plus 4029B (considered as 4029), buccal, lingual, occlusal
aspects respectively.
Ann. S. Afr. Mus., Vol. XLV Plate L
Ss
=
3
scale in cm.
pS Le ee See
a—Hopefield 4025, posterior aspect. b—Hopefield 4026, mesial aspect. c—S.A.M. II715
(Langebaan), astragalus, anterior aspect. d—S.A.M. 11715 (Langebaan), astragalus, posterior
aspect. e—A.M. 19684 (Siwaliks, India). Mandible of Hydaspitherium sp. By courtesy of the
American Museum of Natural History, New York.
Ann. S. Afr. Mus., Vol. XLV Plate LI
cm.
0 5 10 15
St. Arnaud, Algeria: a—1948-1-2, Mus. d’Histoire Naturelle, Paris. Antero-medial aspect of
posterior horn-core of Sivatherium olduvaiense (described by Arambourg, 1948, as Libvtherium
maurusium). b—1948-1-1, Mus. d’Histoire Naturelle, Paris. Medial aspect. (By kind permission
of Professor C. Arambourg.)
Ann. S. Afr. Mus., Vol. XLV Plate LI
a—cast of 1950-1-1 from Lac Ichkeul, Tunis. Buccal aspect. (Original in Mus. d’Histoire
Naturelle, Paris.) 6—cast of posterior horn-core from St. Arnaud (Ain Hanech). (Original
in Mus. d’Histoire Naturelle, Paris.)
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLV
PART V, containing :—
Notes on the Biology of the Lutjanidae (Pisces) of the East African Coast, with
special reference to L. bohar (Forskal). By F. H. Tausot, M.Sc. (With 5
figures in the text.)
SEP 2 196U:sstep zULY 1960 —~PRICE. 3s. 3d.
LIBRARY PRINTED FOR THE
TRUSTEES OF THE SOUTH AFRICAN MUSEUM
BY THE RUSTICA PRESS (PTY.) LIMITED, COURT ROAD, WYNBERG, CAPE
NOTES ON THE BIOLOGY OF THE LUTJANIDAE* (PISCES) OF
THE EAST AFRICAN COAST, WITH SPECIAL REFERENCE TO
L. BOHAR (FORSKAL)
eS a re,
By FP) He Vareor, MSc:
(With 5 figures in the text)
INTRODUCTION
| Lutjanid material for this study has been obtained during routine fishing
from 1954 to 1957 inclusively, by the research ship M.V. Research and her
replacement the M.V. Manihine of the East African Fisheries Research
Organization, Zanzibar, while the author was a member of this Organization.
_ During this period a study of bottom fishes was made, mainly in coral reef
areas of from 3 to 14 fathoms, and also to a lesser extent in deeper water below
the coral reef zone down to 100 fathoms. The work has centred on the reefs
off Lamu on the Kenya coast, in the Mafia Archipelago off the Tanganyika
coast, and on Latham Bank, a shallow bank surrounding a small island
south-west of Zanzibar (see fig. 1). Handlines, gill-nets, trammel-nets, set-lines,
basket-traps, underwater spearing, and explosives have been used for collecting.
In addition fish were occasionally obtained from the local markets on Zanzibar
Island. Information on Lutjanids from the unpublished East African Marine
Fisheries Research Organization records from 1951 to 1953, and for 1958,
has also been used by courtesy of the Director.
This paper is one of a series on hydrographic conditions, Newell (1957,
1959); fish systematics, Morgans (1958), Talbot (1957, 1958), Talbot and
Williams (1956), Williams (1958a, 1959a and b); and fish biology, Talbot and
Newell (1956), Williams (1953, 1956, 1958b), Williams and Newell (1957),
providing some preliminary data on the systematics, distribution and biology
of East Coast fishes of economic importance. A full description of the topography
of the area is given in Williams, 1956. In this paper the systematics of the genus
Pristipomoides is based on Smith (1954) and the systematics of the genus Lutjanus
follows that used in a previous communication (Talbot, 1957).
| The East African coastal area over which this study was made is markedly
affected by the monsoon winds. In all seasons of the year it is bathed by the
* Both the spellings Lutjanus and Lutianus are in current use. The first nomenclatorially
valid use of the generic name is in Bloch, 1790, Nat. ausl. Fische, 4, p.107, in the description of
Lutjanus lutjanus. Cuvier, 1798, Table. elem., Pp- 357 and 705, uses Lutianus (as does Bloch
occasionally after this date), and this is the form used by Jordan and Everman in their Genera
of Fishes, Stanford Univ., 1917, with the footnote “Also spelled Lutjanus’. As Lutianus has not
been universally accepted it seems better to return to the original form Lutjanus.
549
S.A.M. VOL. XLV.
) SMITHSONIAN 411. 2 0 on,
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Chart of the coastal areas where Lutjanids were collected. (From British Admiralty )
Chart 597.) 3
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 551
north-flowing East African Coastal Current, but from May to October
(‘winter’, if one may talk of summer and winter so close to the equator) the
strong south-east monsoon wind speeds up the current to 3-4 knots, and
increases vertical mixing of the upper layers, lowering the thermocline to 50
fathoms. The surface temperature in this period is about 24°C.—25°C. In summer
(November to April) the wind system is reversed, and the moderate north-east
monsoon wind slows up the East African Coastal Current to 1-2 knots. Much
less surfacing mixing takes place, and a very stable surface layer is formed with a
marked thermocline at about 25 fathoms, and a surface temperature of
27°C.—29°C.
Although the majority of species mentioned here are widespread over the
Indo-Pacific region, and many are found from the South African coast up to
the Red Sea and across to Polynesia and the Tuamotu Archipelago, some
10,000 miles to the eastward, apart from systematics little or nothing has been
published about their biology except some recent data on habits and habitats
in systematic papers by American workers aided by U.S. Navy grants (Randall
1955, Schultz 1953, Harry 1953), and the Mauritius Seychelles Fisheries
a Survey undertaken by Wheeler and Ommanney (1953).
The latter authors, in a survey of the coral reef areas lying between
Mauritius and Seychelles, obtained seven species of Lutjanids, all of which
are also found on the East African coastline. Of these seven species four were
seldom caught, but useful data were obtained on the remaining three: L. bohar
(Forskal) (L. civis (C. & V.) of their report), Aprion virescens Valenciennes, and
L. sebae (Cuvier).
In tropical coral areas the catches comprise many more species than
do those of temperate waters, but the numbers of fish of each species are far
smaller. In this survey it was difficult to obtain large enough samples of each
species. With small numbers many established fishery techniques (such as
deductions from length-frequency distribution) cannot be used.
Underwater observations using a Seibe-Gorman aqualung were made on
Latham Bank, about Zanzibar Island, in the Zanzibar channel, and in the
Mafia Archipelago, and showed that two species (L. monostigma and L. vaigiensis)
were common on these reefs although they were very rarely taken by normal
fishing methods. Small samples of these were collected by spear-guns. These
observations also gave a much more accurate picture of relative abundance
of species than did the use of handlines, nets, traps, etc., and some notes on
habits were obtained. It was found that the method was strictly limited in its
usefulness however, for an underwater observer with his trail of bubbles could
not remain hidden, and had a marked effect on the fishes. Many of the smaller
reef species are attracted by the aqualunger, who may become surrounded
by shoals of small fishes. Often also the larger predators such as Plectropomus
maculatus will come closer and inspect the unusual object from mid-water.
Among the Lutjanids these effects can also be seen. Aprion virescens will often
approach close to the diver before continuing down the reef. A shoal of
552 ANNALS OF THE SOUTH AFRICAN MUSEUM
L. gibbus will move off or take shelter. L. bohar after a short time will usually
leave the immediate reef area. Observation is therefore not of a normal
undisturbed reef and its fish fauna—the ‘observer’ has influenced the fish
population, and normal movements and feeding may not be taking place.
This does not imply that the method of underwater observation is not useful.
On the contrary, it is an obvious and developing method which will have
important uses in ecology and ethology (see Reidle 1956). For coral reef
observation however some ‘hide’ method must be used.
That scale rings occur in tropical marine fishes has been shown by a
number of workers (summarized in Menon, 1953). The variation in surface
temperatures from 24°C. to 29°C. between summer and winter in the East
African area might be reasonably considered to be enough environmental
change to affect the formation of annual rings. As is shown below, however,
consideration of L. bohar scale edges with season did not bear this out.
Both scales and otoliths of L. bohar were examined for ringing. The otoliths
showed no clear opaque and translucent zones even on grinding, and although
the surface of the otoliths showed concentric ridges which gave the same counts
as the scale rings in young fishes, in older fishes they were difficult to count due
to crowding towards the periphery, and were also possibly covered over near
the nucleus by further growth. Only the scales are therefore considered here.
Scales of 273 L. bohar were examined, and of these approximately one in four
showed rings considered clear enough to be counted, although rings of more or
less clarity were present in all fish. On an average four scales per fish were
counted, and scales from the same fish with few exceptions showed the same
number of rings at similar relative distances from the scale nucleus. It was
found that individuals of a single sample of fish, taken from the same bank at
the same time did not necessarily have the outermost ring at a similar distance
from the periphery, showing that the rings were not all formed at the same
time of year in all fish. This suggests that the rings that do form are due to
spawning, and not to seasonal changes. With checks forming at different times
of year with different fish it is impossible to test whether the rings are annual
by the method of watching the periphery of the scales of samples of fish
periodically during the year.
Forty scales were re-read a year after first reading to check error. Of these
19 were re-read exactly as in the first reading, 12 disagreed by one year, and
9 by more than one year. These discrepancies are due to faint rings being either
considered as false checks or true annuli. This error can be stated thus: in
about 47°% of scales used here reading error is negligible; and in about 78% of
scales an error of + 1 ring may be present. If it is realized that the scales
being read here are already only those showing the clearest rings (one-quarter
of the total) it is obvious that the clarity of ringing does not approach that of
many temperate species. Clark (1958) for example in the re-reading of young
haddock samples by the same worker obtained 90% and 93% similarity.
Nevertheless in many specimens of L. bohar consistent ringing is present in the
ee a a
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 553
scale structure, due to some regular change in the metabolism of the fishes.
It is possible that these periodic changes are not annual, but their regularity
suggests that they will be found to be connected with the spawning cycle or
be due to periodic feeding changes. As has been suggested above, the former is
the more likely answer. Wheeler (in Wheeler and Ommanney, 1953) has
suggested that L. bohar spawns twice a year. As will be seen in this report the
results from East African coastal fish rather suggest an extended breeding
season in the warmer months. Lacking evidence to the contrary the ringing
here seen is considered as annual.
DESCRIPTION OF GONAD CONDITION
Gonads were described macroscopically in the fresh condition as the
fish were gutted on board. For females a system based on seven stages was
used (after Bowers, 1954). For males five stages were discernible.
Males: I. Immature. Gonad small, usually threadlike, no sperm extruded
on cutting and squeezing.
II. Mature unripe. Gonad small, sperm extruded on cutting and
squeezing.
III. Ripe. Gonad enlarged and full of sperm.
IV. Ripe running. As above, but milt extruded on pressure to flank.
V. Spent. Testis shrunken, not full and round, little sperm.
Females: I. Jmmature. Ovaries small and threadlike, eggs microscopic.
II. Mature unripe, or Virgin maturing. Gonad of moderate size, eggs
microscopic, gonad often translucent. The two stages may
sometimes be distinguished as there may be remains of corpora
lutea in the mature fish visible as small orange flecks in the
ovary.
III. Mature ripening. Ovary of moderate size, eggs visible to the
naked eye, opaque.
IV. Nearly ripe. Ovary enlarged and extended, eggs clearly visible,
opaque.
V. Ripe. Ovary enlarged and distended, tunica breaks easily, some
eggs transparent.
VI. Ripe running. Nearly all eggs transparent, eggs extrude on slight
pressure to flank.
VII. Spent. Ovary flaccid, shrunken, and with some residual eggs.
Although stage II males are called mature unripe it seems probable that
sperm may be present in the testis before the fish are capable of mating, as in
L. bohar a very small gonad holding some sperm may be found in fish of about
554 ANNALS OF THE SOUTH AFRICAN MUSEUM
200 mm. but the first males found with enlarged ripe gonads were of a much
greater size than this. State II contains both virgin developing males, and
mature males in resting condition. No macroscopic difference was noticed
between the two.
Species No. Examin. Immature Mature
Lutjanus bohar (Forskal) 854 443 (150-439 mm.) 411 (440-660 mm.)
Aprion virescens Valenciennes 259 18 (202-452 mm.) 241 (460-800 mm.)
Lutjanus rivulatus (Cuvier) 129 4 (395-449 mm.) 125 (450-640 mm.)
L. fulviflamma (Forskal) 126 41 (51-159 mm.) 85 (160-220 mm.)
L. gibbus (Forskal) 121 10 (170-219 mm.) III (220-355 mm.)
L. sanguineus (Cuvier) 102 39 (170-479 mm.) 63 (480-650 mm.)
L. kasmira (Forskal) aa — 77 (125-205 mm.)
L. sebae (Cuvier) 27 12 (128-489 mm.) 15 (490-665 mm.)
L. monostigma (Cuvier) 18 7 (275-349 mm.) I1 (350-420 mm.)
L. argentimaculatus (Forskal) 13 9 (300-459 mm.) 4 (460-630 mm.)
Size Range
Pristipomoides microlepsis Bleeker 12 260-640 mm.
Lutjanus ehrenbergi (Peters) 6 44-98 mm.
L. vaigiensis (Quoy and Gaimard) 6 200-250 mm.
L. lineolatus (Ruppell) 5 120-175 mm.
Pristipomoides typus (Bleeker) I 525 mm.
Aphareus rutilans Cuvier I 780 mm.
Lutjanus duodecimlineatus I 150 mm.
TasB.e I
A list of the Lutjanids taken during the survey, with the proportions of mature
to immature fish. (For the commoner species the smallest length at which
mature fish were found is used for a division into ‘mature’ and ‘immature’
fishes in the table.)
Lutjanus bohar (Forskal)
L. bohar is one of the commonest predators of exposed coral reefs in the
East African coastal area. The majority of specimens were taken by handline,
but the species was also taken in basket traps, trammel nets, and very
occasionally on trolled lures. It is fairly common in local markets, seldom in
abundance, but present in regular quantities throughout the year. Wheeler
and Ommanney (1953) took larger numbers and a greater total weight of this
species than any other on the Mauritius-Seychelles banks.
This species sometimes causes ciguatera poisoning in the Mauritius area
and is banned in the markets there. Harry (1953) states that in Raroia Atoll
of the Tuamotu Archipelago ‘large adults of L. bohar are poisonous, and natives
know to a few inches of length when an individual is poisonous or not’. Randall
(1958) in his review of ciguatera mentions this species as causing poisoning
in a number of areas. Whitley (1943) in his list of poisonous fishes of Australia
includes the closely allied (perhaps synonymous) L. coatsi. In spite of this there
is no record known to the author of this species being considered poisonous in
the East African coastal region, and both there and in the Seychelles (Wheeler
and Ommanney, 1953) L. bohar is considered a prime market species. Randall’s
suggestion that this type of poison enters the fish through one of its foods,
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 555
probably a blue-green alga, is consistent with the species being poisonous in
some areas and not in others.
Handlining showed this species to be common on exposed areas with
actively growing coral in from 4 to 15 fathoms. Adults were seen underwater
on the outer slope of the fringing reef, and entering into gaps in the fringing
reef where these were deep (five to seven fathoms), but were not seen in areas
where the channel was shallower than this, or on the reef flat at high tide,
(see figs. 4 and 5). Juveniles of about 100 mm. were seen in and about coral
where the fringing reef channel was under 5 fathoms deep. On the outer edge of
Tutia Reef and on Latham Bank the species was usually seen in loose and
actively moving shoals of about two to seven fish, over coral in mid-water
in 3 to 10 fathoms. Underwater observation did not go deeper than this.
Shoals were loosely knit, and seemed to break up and rejoin, with no tight
cohesion as with many other species of the genus. What shoaling instinct there
was, however, results in the species being more often seen in twos and threes
than singly. Unlike many related species which may stay around one coral head
for the time observed (maximum 2 hours), L. bohar is continually on the move,
as though actively searching for food over wide areas. Actual feeding has not
been observed underwater.
L. bohar was also found to be common in deep water of the Kenya coast in
the region of Lamu (see fig. 1). In this area the continental shelf is wider than
in most of the East African coastal region, and rich populations of fishes were
found on grounds approximately 25 to 65 fathoms deep. L. bohar was caught
from 25 to 46 fathoms, but where a bathythermograph was used while fishing
it was never found below the major thermocline, which varies from 25-50
fathoms with season (Newell, 1957). The dominant species in these rich
populations were Lutjanus bohar, Lutjanus rivulatus, Epinephelus undulosus, Lutjanus
sanguineus, Lethrinus waigiensis, and Lethrinus kallopterus, in order of abundance
(Williams, 1958).
Sampling showed that this species is present on the reefs at all times of
year and showed no evidence of migrations. Underwater observations proved
that on certain small reefs the species could be present at one visit and not
at the next.
FEEDING
Of the 854. L. bohar examined, 58% of the fish caught had empty stomachs
or contained only bait. Table II lists the food organisms found in L. bohar
stomachs. The species is a euryphagous predator, feeding basically on fish
(see fig. 2), but capable of eating crawfish, crab, prawn, squid, octopus,
ophiuroids and even pelagic pteropods, doliolids and pyrosomas when these
are abundant in the plankton. Although a wide variety of small reef fishes was
the main food over the period of study, at certain times when there was an
abundance of any one organism the samples contained nothing but these
organisms. This was clearly seen in two samples from North Mafia Bay
556 ANNALS OF THE SOUTH AFRICAN MUSEUM
(Cumulus Bank), and once in a deep-water sample from Lamu. In December
1952 a large proportion of the samples from these two banks was packed with
Penaeid prawns, probably migrating to or from the mangrove areas of the
huge Rufiji delta. In November 1953 there was an unusually large amount of
a larval Stomatopod in the Mafia-Latham Island area. These were seen
swimming on the surface at night, and almost everything caught from bottom-
dwelling Epinephelids to pelagic Sphyraenids contained them. All Lutjanus
Fic. 2
Relative abundance of food organisms found in Lutjanus bohar.
bohar from Vulture, Cumulus and Latham Banks were full of Stomatopod
remains. In one particular sample (14-19 November 1956) of the many taken
from deep water off Lamu, L. bohar, L. sanguineus, L. rivulatus, Aprion virescens and
one species of Ephinephelus all contained salp tests. Otherwise salps were not a
common item of food in any of these species. L. bohar then follows the general
pattern that most predatory species at any one time will eat the commonest
foods available to them in their particular habitats (Stephen 1930, Allee et
al. 1949).
Some change of diet with size was shown. The smallest L. bohar taken
(150 mm.) were already fish predators and fish was the predominant food
throughout the size range investigated. Crabs were first found in fish over
200 mm. in length, and Cephalopods only in fish over 250 mm. As the fish
ea ee ee
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 557
increased in size above this Cephalopods became an increasingly important
food.
Wheeler (in Wheeler and Ommanney, 1953) found essentially similar
feeding for L. bohar of the Mauritius-Seychelles banks, although a greater
frequency of crustacean and plankton food was found (Fish 114, Crustaceans
117, Plankton 116, Molluscs 83, other 31). Wheeler also concluded that
plankton is taken only in times of special density.
Reptilia Crustacea
Green Turtle (Chelone midas L.) juvenile. | Charybdis natator (Herbst)
Calappa sp.
Pisces Charybdis sp.
Carangid fish Lupa sanguinolenta (Herbst)
Holocentrus sp. Achelous sp.
Ostracion sp. juvenile Thalamita sp.
Echidnid eel. Monomia sp.
Scaridae (many unident. species) Xanthid crab
Lethrinus chaerorhynchus Oxyrhynchid crab
Lethrinus latifrons Ruppell Panulirus sp.
Syngnathid fish Penaeus sp.
Canthidermis sp. Pagurus sp.
Monacanthid fish juvenile Metapenaeus sp.
Clupeidae Scyllarid larva
Mullidae. Megalopa larvae
Sphaeromid Isopod
Tunicata Amphipods
Pyrosoma Stomatopod
Doliolid. Stalked cirripede.
Echinodermata Polychaeta
Ophiuroids. Polychaete bristles.
Mollusca Plants
Cavolinia sp. Cymodocea leaves
Turbo sp. (‘Green Snail’) Green alga.
Octopus
Squid
Tectibranch rem.
Tase II
Food organisms found in Lutjanus bohar.
GROWTH RATE
Wheeler (op. cit.) has estimated the growth rate of L. bohar on the Chagos
Bank (Seychelles) from length-frequency curves, and has suggested that the
one-year group is 180 mm. total length (136 mm. standard length), with
additions of 120 mm. and 110 mm. in the second and third years respectively.
He estimated that on the Seychelles plateau and the Amirantes 510 mm.
total length (425 mm. standard length) was attained in four years. The Peterson
method of age determination is dependent on a short spawning period and
roughly equal growth rate of the individuals of each spawning. Wheeler found,
in over 2,000 fishes taken during 1948-9, that ripe females were present only
in October and November and again in March, suggesting that the first of
these tenets is satisfied, with due allowance made for the addition of a new
group every six months and not every year. As different banks were considered
558 ANNALS OF THE SOUTH AFRICAN MUSEUM —
separately there is every reason to suppose that growth rate is approximately
the same and that the method is valid for L. bohar in the Seychelles-Mauritius
area. In the East African coastal region, however, the breeding season seems to
be an extended one, ripe female fish being taken over most of the year. This
and the small size of samples has precluded the use of the Peterson method here.
Wide, diffuse rings (formed of a group of fine rings) were present in
juveniles, but in adults the scale checks were often sharp, with the lamellae
after the check beginning at different angles, as though either resorption had
taken place, or growth had begun again after complete cessation. If the checks
in the adults are spawning checks, the diffuse rings in the juveniles are possibly
due to a physiological sex rhythm already existing in the juvenile, as suggested
for the Hake by Hickling (1933).
Peripheral checks were seen in September, November, January, February,
March, April, and May. This is mainly in the north-east monsoon period, which
lasts from November to April, when the water temperatures are higher than
the period of the strong south-east monsoon from May to October. The largest
L. bohar taken during this survey and aged
St. Length to Annual was 660 mm. standard length, and showed 12
Age nearest5 mm. Increment : ‘
as , ; growth rings. On one fish of 615 mm. 13 rings
3 240 i were found. The smallest fish whose scales
: ae a were read was 200 mm., and showed three
6 420 50 growth checks. Increments were 70 mm. and
: ae - 60 mm. in the fourth and fifth year, and then
9 550 40 gradually reduced to approximately 35 mm.
oy 385 oe per year (see Table III). Maturity was reached
about the sixth to seventh year (450 mm.) and
Tasre Iil no difference in growth rate was seen between
Average lengths and the sexes. 7
increments for different This is a very much slower eo. rate
age groups of L. bohar. ‘than that given by Wheeler, increments being
Maturity reached at 6-7 about half as much from scale ringing. The
years. possibility cannot be excluded that two rings
are laid down per year, although at present
there seems no obvious reason for such double ringing in the coastal fishes.
MATURITY AND SPAWNING
No distinct spawning periods were found in L. bohar. Few ripe (Stage V)
females were found, these being taken in July, September, November, February
and March. This suggests an extended breeding season over most of the year. ©
The double spawning period suggested by Wheeler (op. cit.) for L. bohar in
the Seychelles area is not proved for coastal fish by these results. The smallest
females with enlarged gonads (recorded as ripe or nearly ripe) were 445 mm.
Although sperm was seen in males as small as 270 mm., when the testes are
still threadlike, the first ripe (Stage III) males were found at 450 mm.
ea oo,
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 559
Aprion virescens Valenciennes
Aprion virescens is a fast-moving predator common over coral areas in 3 to
15 fathoms, feeding from the surface through the mid-water region to the
bottom. It is the only Lutianid species to be regularly taken by surface lures.
259 specimens were taken by handline and surface lure from 202 mm. to 800
mm. standard length (25 lb.). This species was often taken when handlining
for L. bohar, usually on non-weighted lines. Over coral reefs in from 4 to 14
Fic. 3
Relative abundance of food organisms found in Aprion virescens.
fathoms L. bohar is typically caught from mid-water to the bottom, and Aprion
virescens from the surface to mid-water. They therefore occupy distinctly
different, but overlapping habitats. A. virescens has been caught on handlines
fishing down to 50 fathoms. The statement by Williams (1956 p. 37) of this
species being taken on lines fishing from 75-80 fathoms is an error, and refers
to a specimen taken fishing at 50 fathoms off the north end of Pemba Island on
23 October 1953. More than once this species has followed handlines being
hauled in from deep water (50-60 fathoms) to the surface. It is possible that
the species, which is mainly known as pelagic, may take the bait as the lines
are being hauled in, and not at the bottom. There is no proof at present of the
species occurring below the major thermocline in the colder sub-surface water.
A. virescens is considered a prime food fish, and occurs in the local markets
throughout the year in small numbers.
Underwater, A. virescens is commonly seen in coral areas (Mafia, Latham,
Cumulus, Zanzibar Channel), always actively moving in mid-water, never
560 ANNALS OF THE SOUTH AFRICAN MUSEUM
sheltering in coral. It is usually in loose and widely spaced shoals of two to
five fish, but occasionally solitary.
FEEDING
A. virescens seems able to feed from the surface to the bottom. Fish was the
most important food taken, and comprised Lethrinus microdon, Iniistius sp.,
Siganids, Tetraodonts, Scarids, Balistids, Labrids, Synodontids and Atherinids.
Plankton was also often found in the stomachs of even the largest fishes, and
included fish eggs, larval fish, stomatopod larvae, salps and zoeae larvae.
Crustaceans, mainly Portunid crabs, and to a lesser extent Penaeid prawns,
were also important. Squid was occasionally taken. (See fig. 3.)
MATURITY AND SPAWNING
The smallest female recorded as ripe (Stage V) was 465 mm. standard
length, and another female of the same length as mature ripening (State IV).
Gonads of males were seen with sperm at 410, 420, 455 and 460 mm., but the
smallest males recorded with enlarged full testes were just under 500 mm.
Ripe females were found only in December, January and February,
suggesting a breeding season during the warmer water of the north-east
monsoon period. Nearly ripe fish (Stage IV) were recorded in most months,
and more information will probably prove an irregular extended breeding
season.
Lutjanus rivulatus (Cuvier)
One hundred and twenty-nine specimens were taken by handline and
underwater spearing, ranging from 395-640 mm. (maximum weight of 19 lb.).
This species had rarely been taken from the E.A.M.F.R.O. research vessels
until the rich fish populations at Lamu off the Kenya coast had been found
(mentioned under L. bohar) in 25-65 fathoms. L. rivulatus formed about 20%
of the catch in these areas. Underwater observation has shown that although
seldom caught by handline over shallow coral reefs, L. rivulatus is common in
certain sheltered coral areas such as Tutia Gap (Mafia) and inside the fringing
reef in 5-7 fathoms with L. bohar on the east coast of Zanzibar Island, and also
at Ras Kizimkazi at the southern tip of Zanzibar Island. At these places
occasional mid-water shoals of five to ten fish, often in conjunction with
L. bohar and L. argentimaculatus are present. The species has also been seen
singly sheltering under dense beds of the ‘platform coral’ (Acropora hyacinthus
[Dana]) on Tutia Reef, and in rocky areas at 5 fathoms on Latham Bank.
FEEDING
L. rivulatus is predominantly a fish predator, also taking crabs, polychaetes,
squid, octopus, echinoids, ascidians and polyzoa. It is chiefly a bottom feeder.
One sample from deep water contained many fishes filled with salps.
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 561
MATURITY AND SPAWNING
Maturity is reached in both males and females at about 450 mm. standard
length. Ripe females were found in February, March, April, November and
December, and ripe males in March, April, August, November and December,
suggesting an extended breeding season in the warm north-east monsoon
period.
Lutjanus fulviflamma (Forskal)
This species is abundant over the whole East African coastline extending
as far south as South Africa (33°S.), and although small in size is an important
species economically, always present in the local fish markets, often in large
numbers. It is common in the fringing reef channels, the outer reef slope,
mangrove areas, the reefs of Zanzibar channel, the Mafia area and in estuaries
(see figs. 4 and 5). Juveniles have been seen in pools on the reef flat, and in
shallow water of from six inches to a foot around Zanzibar town over both
sandy and weedy bottoms. No other Lutjanid species has as wide a distribution
of habitats as L. fulviflamma.
One hundred and twenty-six specimens were examined, taken by basket-
trap, handlines, trammel-nets, and bought from the local markets. The size
range was from 51-220 mm.
FEEDING
Crustaceans were the predominant foods; being mainly crabs (including
Portunids and Callapids) and also Eupagurids, Sphaeromid isopods, Penaeid
prawns and Stomatopods. Fish remains included Engraulids, Fistularids and
gobies.
Different samples often contained foods of one type, presumably indicating
local abundance of one particular food organism in the area in which the
sample was obtained. Most of the food organisms found were bottom animals.
The investigation of small samples of this species in Durban Bay (South
Africa) by the Zoology Department, University of Cape Town, has shown
very similar results. Bottom-dwelling crustaceans (mainly Hymnosoma orbiculare
Dem. and Penaeid prawns) predominated in the stomachs, with fish (Eleotrids,
gobies and Lutjanus sp.) next in importance (from unpublished records by
courtesy of Professor J. H. Day).
MATURITY AND SPAWNING
Of 112 fishes whose gonads were examined, 36 were immature and
unsexed, of the remaining 76, 51 were female and 25 male. Of one sample of
13 fish, 12 were female and 1 male. Males were recorded with sperm as small
as 145 mm., and the first fully ripe male was found at 170 mm. Ripe females
were recorded as small as 160 mm.
ANNALS OF THE SOUTH AFRICAN MUSEUM
562
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564. ANNALS OF THE SOUTH AFRICAN MUSEUM
Ripe females were found in March, August, October and December,
suggesting an extended breeding season mainly in the north-east monsoon
period.
Lutjanus gibbus (Forskal)
Lutjanus gibbus is a small species, seldom reaching more than 4 lb. in weight,
occasionally seen in local fish markets, but seldom in large numbers.
One hundred and twenty-one specimens were taken by handlining and
trammel-nets, always on the bottom and never in mid-water with L. bohar
and A. virescens. The size range of fish taken was from 170 to 355 mm. This
species was usually taken at night. It was only found in shallow water of from
3 to 8 fathoms.
Underwater observations showed that this species keeps a few inches
above the coral, often sheltering in the branches of ‘stag coral’ (Acropora
formosa [Dana]), and in the leaves of ‘platform coral’ (Acropora hyacinthus
[Dana]). It was seldom seen singly but usually in dense, closely knit shoals,
typically roving over the bottom in a single layer, closely following the bottom
contours. Numbers ranging from one to fifty fish and over were seen, but usually
shoals were from ten to twenty-five. The species is common on exposed and
sheltered coral reefs with rich coral growth, and was seen at Tutia, Vulture,
and Cumulus banks in the Mafia area, and on the fringing reef outer slope at
Zanzibar. It was also taken on Latham bank and at Lamu. Adults were never
seen in the fringing reef channels. Juveniles of about 50 mm. long were seen
on the reef flat, and are occasionally taken in beach seine hauls on Zanzibar
Island.
FEEDING
Foods eaten were mainly crustaceans, including crabs and Penaeid
prawns. Small coral fishes were also occasionally taken. Coral and sand were
sometimes present in small quantities.
MATURITY AND SPAWNING
The smallest mature females with gonads approaching spawning condition
(Stage IV) were found at 223, 235, 240 and 245 mm. standard length. The
smallest mature males were 240 and 280 mm. Ripe fishes (either sex) were
found in March, November, September and December, i.e. the north-east
monsoon period.
Lutjanus sanguineus (Cuvier and Valenciennes)
One hundred and two specimens were taken by handline and basket-trap,
ranging from 170-650 mm. (13 lb.). Juveniles were taken in basket-traps in
the Zanzibar channel and in the Mafia area, in 6 to 7 fathoms on coral and
;
;
|
2
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 565
Cymodocea bottoms, and were also found in the Chukwani fish-ponds (Zanzibar
_ Island), a mangrove area. Adults were not taken by basket-trap, or by hand-
lining in daylight over coral reefs. On one bank in the Mafia area (Snapper
Knoll, Niororo Island) adults were taken on four occasions by handlining at
night. Adults and juveniles were also taken by trammel-nets overnight in
shallow water in Lamu Harbour. Adults of Z. sanguineus were common in
_ 25-47 fathoms off Lamu, and were one of the dominant species in these deep-
water catches (mentioned on p. 555).
At certain times of year (January, February and March) this species
occasionally floods local markets (Zanzibar Island), being taken off the southern
tip of Zanzibar Island in 40 fathoms. Off Shimoni (Kenya) it is also taken in
quantity at certain times of the north-east monsoon (November to April).
L. sanguineus was never observed underwater.
When it is caught it is seldom taken singly, but usually a number within
a few minutes suggesting a shoaling habit.
FEEDING
Fishes were the commonest food (including Syngnathus biaculeatus,
Monacanthids and Apogonids) but Penaeid prawns, crabs, stomatopods,
cephalopods and plankton (salps, doliolids, pteropods and medusae) were also
found. No change in diet with size was found.
MAtTuRITY AND SPAWNING
The smallest males and females with ripe gonads were 480 mm. and
505 mm. respectively. Stage IV females were found in March, August,
September and November and Stage V females in April and August. Stage
III males were found in March, April, August and November.
Lutjanus kasmira (Forskal)
Seventy-seven specimens were caught by trammel-net, underwater
spearing, and handline, ranging from 125-205 mm. L. kasmira is a small species
seldom reaching 3 lb. in weight, and is not caught unless very fine lines and
hooks are used.
Underwater it is seen to be abundant, often in dense shoals of 25 fish and
more, never singly, and usually about actively growing coral in exposed
areas. It has been seen off Ras Nungwe (Zanzibar), Tutia Reef, and outside
the fringing reef on the Zanzibar east coast, in 2-5 fathoms. One specimen
taken in deep water (35 fathoms) off Malindi (Kenya) differs slightly in
coloration, scaling, and the number of dorsal spines from the shallow water
specimens, and is probably a deep water race. This specimen has been described
in a previous communication (Talbot, 1957). The species is not found in
sheltered mangrove areas, and has not been seen about the Zanzibar channel
reefs. It is occasionally seen in local markets.
566 ANNALS OF THE SOUTH AFRICAN MUSEUM
FEEDING
Crustaceans were the predominant food and included crabs and
amphipods. Squid, fish remains and algae were also found.
MATURITY AND SPAWNING
Females were mature at the smallest sizes taken, i.e. 125 mm. Males were
first seen with sperm at 155 mm., and first recorded as ripe at 165 mm.
Ripe fishes were found in March and November suggesting pies in
the warm water north-east monsoon period. |
Lutjanus sebae (Cuvier)
Twenty-seven specimens were taken by handlines (adults), basket-traps
and spear-guns (immature fish), ranging from 128 to 665 mm.
The juveniles of this species occur in shallow water (5-10 fathoms) and
fish up to 360 mm. (34 lb.) have been taken on banks in the Mafia archipelago
(Snapper: Knoll near Niororo Island and Sefo Reef). Larger specimens have
been found to be fairly common in deeper water of 20-48 fathoms and were
taken off Tutia Reef and on the Lanu Banks. Occasional specimens of up to
60 lb. have been seen at Ras Kizimkazi (Zanzibar Island) taken on handlines
in 40 fathoms by local fishermen. L. sebae occurs regularly in small quantities
in Zanzibar markets. It was never seen underwater by the author but has
been seen in 5 fathoms on Sefo Reef by Dr. J. F. C. Morgans (personal
communication).
FEEDING
Stomachs contained fish, stomatopods, crab and cephalopod remains.
MATURITY AND SPAWNING
Females with developed gonads (Stages IV and V) were only found above
490 mm. standard length. Insufficient data were obtained to estimate size at
which males mature. Breeding is in the north-east monsoon period, ripe fish
having been found from November to March.
Lutjanus monostigma (Cuvier)
Eighteen specimens were taken, 15 by underwater fishing and 3 by hand-
lining in 1-4 fathoms. Size range was from 275 to 420 mm. This species is very
rare in the local markets. Underwater observation, however, showed it to be
common in areas where large coral growths form deep shelter, and it seems
completely limited to this type of habitat. It was common about Tutia Reef,
Latham Island, in the Zanzibar Channel, and was seen occasionally inside
the fringing reef on the Zanzibar east coast. It was never seen in shoals although
;
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 567
two or three were often seen under one coral shelter. Individuals often remained
under one coral for the whole period of observation (up to two hours).
FEEDING
Fish remains (including one Mullid and one Labrid) were present in
most stomachs, and Penaeid prawn remains were also found.
MATuRITY AND SPAWNING
Ripe or nearly ripe females (Stages [V—V) of 395 mm., 390 mm., 420 mm.,
and 400 mm. were taken. No ripe males were caught. From the meagre
data the fish appear to mature at over 350 mm. (2 lb.).
Ripe females were found in November and February.
Lutjanus argentimaculatus (F orskal)
iEhistcen specimens of 300 to 630 mm. (15% lb.) were taken from abeléewed
reef areas in up to 7 fathoms by handlining or set nets at night. This species is
abundant in East African coastal waters, and is an important market species.
It does not. occur on exposed coral reefs, however, and is therefore not well
represented in the E.A.M.F.R.O. catches. It is very common in. shallow
mangrove areas and estuaries, and common in sheltered waters such as the
Zanzibar channel. It was commonly seen during underwater observation
inside the fringing reef off the Zanzibar east coast, in the semi-estuarine waters
of Chwaka creek (Zanzibar) and about large, and often dead, coral growths in
the Zanzibar channel. It may occur singly or in shoals of up to 20 fish. Juveniles
are fairly common in sheltered mangrove areas. |
L. argentimaculatus was never seen during underwater observation on
exposed coral reefs.
MATuRITY AND SPAWNING
Males were found with testes containing sperm at 330 mm., 410 mm.
and 460 mm., and one ripe male of 515 mm. was taken in November 1958.
One female was considered mature at 460 mm., and one De female of 630 mm..
was taken in November 1957. |
Lutjanus ehrenbergi (Peters)
Six specimens were taken from mangrove pools on Zanzibar Island
(Chukwani fish-ponds) from 44 to 98 mm. standard length. Two of the.
specimens were fully mature females of 75 mm. These are the smallest mature
Lutjanids found during the survey. .L. fulviflamma, also maturing at a relatively
small size, was first found mature at 150 mm. L. ehrenbergi was never seen during
underwater observation.
568 ANNALS OF THE SOUTH AFRICAN MUSEUM
Lutjanus lineolatus (Ruppell)
Five individuals of this small species were taken (standard lengths
120-175 mm.) but no biological data were obtained from them. Underwater
observation showed this species to be often present in large shoals (30 to over
100 fishes) on exposed coral reefs, often in conjunction with L. kasmira. It is
also fairly common in East African markets, and its rarity in E.A.M.F.R.O.
catches are due to catching methods.
Lutjanus vaigiensis (Quoy and Gaimard)
Six specimens were taken by underwater spearing and handlines from
200 to 250 mm.
This small species has very occasionally been seen in local markets, and
underwater observations shows that it is present, although not common, on
shallow coral reefs (Ras Nungwe, Tutia Reef, inside the Zanzibar fringing reef),
usually singly, but occasionally in pairs. It is always in or near coral shelter.
It is often present in the same areas as L. monostigma. Shoaling in this species
has never been seen. Juveniles have been taken in a mangrove area (Chukwani
fish ponds, Zanzibar Island). Fully mature males were found at 190 and
205 mm.
Pristipomoides microlepis (Bleeker)
Twelve specimens were obtained by deep lining (47-67 fathoms) ranging
from 260 to 645 mm. (14 Ib.).
This species has been taken off Malindi, Pemba, and Tutia Reef. Smith
(1954) records it as a major component of the Shimoni (Kenya) deep-water
fishery. The species lives in the cold sub-surface water below the thermocline,
and has not been taken in shallow water.
Pristipomoides typus (Bleeker)
One specimen of 525 mm. (84 Ib.) was taken off Tutia Reef in deep water
(about 55 fathoms). Smith (1954) records it as occurring in the deep-water
Shimoni fishery.
Aphareus rutilans Cuvier
One specimen, a mature female of 785 mm., was taken at 60 fathoms off
Lamu. Water temperature at that depth was 19°C. from a bathythermograph
reading.
Lutjanus duodecimlineatus (Valenciennes)
One specimen was purchased in the Zanzibar fish market (150 mm.).
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 569
CONCLUSION
Few Lutjanids are restricted to the East African coast. Of the seventeen
species recorded here, all except two (L. ehrenbergi and L. duodecimlineatus)
reach the Australo-Pacific region. In the reverse direction distribution of this
family is not so uniform, however. Many Lutjanids found in the Parific Ocean
and the Eastern Indian Ocean are not found in the Western Indian Ocean.
For example, in the genus Lutjanus 32 species occur in the Hawaiian Islands
(Herre, 1953), excluding the freshwater LZ. maxwebert and the doubtful
L. philippinus. Of these only 13 are found in East Africa. This suggests a centre
of origin, or at least strong adaptive radiation in the Australo-Pacific region.
At present there is clearly difficulty of dispersal in an east-west direction for
many species, but whether this is due to paucity of suitable environments in
the north and western Indian Ocean, or to some physical barrier to migration
is not obvious. Temperature, which limits the southerly distribution of this
typically warm-water family down the coast of Africa, does not operate as a
barrier to its spread along the Indian and Iran coasts. Lutjanid distribution
bears out Ekman’s statement that ‘the rich Indo-Malayan fauna is distributed
over a large part of the Indian Ocean, but the number of species constantly
decreases as we proceed in a westerly direction’.
Down the African coast some species disappear at Delagoa Bay (26°S.),
with the last coral reefs, and then there is a steady decrease in species to East
London (33°S.), no members of the genus Lutjanus, and only the genera Aprion
and Etelis (Indo-Pacific, not recorded during this survey) reaching farther south,
to Knysna (34° 5/S.). (Note: Two specimens of L. sanguineus have been taken
in Algoa Bay, and one at Plettenberg Bay, 34°S., during 1958-9.) The distribu-
tion of this family is paralleled by that of the reef-building corals. Although
no coral reef growths have been found south of Delagoa Bay, reef-building
genera are found to just north of East London (Stephenson, 1947). Coastal
temperatures drop rapidly between Port St. Johns (31° 40'S.) and East
London (33°S.) due to an outward turning of the Agulhas current. The latter
port coincides with the 20°C. isotherm in winter (Sverdrup et al., 1942).
Vertical distribution is very limited in most of the family (see fig. 4).
Of the seventeen species taken eight species were never caught deeper than 14
fathoms. Of these most were common about coral reefs, and their deepest
limits coincided with the limit of vigorous coral growth, which was usually
between 10 and 15 fathoms. It is possible that the same factors might be the
cause of both the limit of distribution of the fishes and the end of reef growth,
but more likely that the fishes are limited to the coral habitat.
The thermocline, varying from 25-50 fathoms, is deeper than the foot of
the actively growing coral area, and at 10-15 fathoms water conditions differ
little from the surface. Temperatures at this depth were not found below 24°C.
by Newell (1957) and in summer are very much above this, so temperature
does not seem to be a barrier in this connection. Suggested reasons for the
downward limit of coral growth will be discussed in a later paper; here it will
570 ANNALS OF THE SOUTH AFRICAN MUSEUM
be sufficient to state that for L. vaigiensis, L. monostigma, L. lineolatus, L. gibbus,
L. fuloiflamma and the shallow-water form of L. kasmira (referred to on p. 565),
the foot of the living coral reef (excluding tallus slopes) is the downward limit.
Although some physical environmental factor or combination of factors may
be the cause of this, the abrupt ending coincident with that of the coral suggests
that in a downward direction the fish distribution is determined by the latter.
L. fulviflamma was not restricted to coral areas but was also abundant in
sheltered areas such as mangrove swamp channels and Cymodocea beds. L.
argentimaculatus, also limited to shallow water, was found in more sheltered
habitats only, including mangrove areas, the boat channel of fringing reefs,
and sheltered coral. L. ehrenbergi was taken only in a mangrove swamp.
L. bohar and L. rivulatus were for a long period (1951-5) considered to have
only a shallow-water distribution, never being taken below about 14 fathoms.
In 1956, however, when the deep-water (25-65 fathoms) banks off Lamu,
Kenya, were fished it was discovered that these two species were often abundant
at much greater depths, reaching 46 and 51 fathoms respectively. In these
areas the shelf is unusually wide for the East African coast (considered by
Morgans, 1959, to be due to the deposition from an old river delta), and rich
feeding-grounds are present. It seems clear that this favourable habitat is the
reason for the presence of L. bohar and L. rivulatus in deeper water.
On these banks the bottom temperature may vary from 22°C. to 29°C.
Newell also gives one reading of 18°C. at 50 fathoms. (Newell 1959, Morgans
in unpublished E.A.M.F.R.O. reports.) From the foods taken by these members
of the genus Lutjanus it appears that they are all bottom feeders, and it is
probable that temperature becomes an important factor in the distribution of
these members of this genus to the colder deeper portions of these banks, if
we may judge by their distribution down Africa, referred to above. It is quite
clear from the great deal of fishing that has now been done in this area that the
genus Lutjanus is only found in water of the East African coastal current, and
not below the thermocline. No members of this genus have been taken in
water below 23°C. where a bathythermograph has been used in conjunction
with fishing. In addition to L. bohar and L. rivulatus this also applies to L. sebae
and L. sanguineus whose adults were common on these banks. In the former only
juveniles and young fishes up to 360 mm. were taken in shallow water, and
adults in water of 30-49 fathoms. Adults of the latter extended from the shallow
water of Lamu Harbour (a mangrove area of 4 fathoms) to 47 fathoms, and
were more abundant in deep water. Juveniles of these species were only taken
in sheltered water.
In contrast to the genus Lutjanus the two species of the genus Pristipomozdes
and Aphareus rutilans were taken in 47-67 fathoms and 60 fathoms respectively
and never in shallower water. Also found on the Lamu Banks, these species are
usually present below the thermocline. A. furcatus, not taken during this survey
but recorded by Smith (1954) in East Africa, seems to be a shallow-water
species, being referred to by Randall (1955) as fairly commonly seen underwater
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 571
in shallow coral-rich areas of the Gilbert Islands, and taken on the surface on
lures by Schultz (1953) on Bikini Atoll. Smith’s record is of one specimen from
20 fathoms off Pemba Island.
The genus Aprion has a pelagic habit and a depth range to about 50
fathoms, but is more abundant in shallow water over the reefs.
Clear differences were found between the catches of Lutjanids from coral
areas facing the open Indian Ocean and exposed to violerit wave action, and
those of sheltered coral islands in the 15-mile channel between the African
mainland and Zanzibar Island or from the inner Mafia Archipelago (see fig. 1).
L. argentimaculatus was taken both in the boat channel and also from sheltered
coral reefs, but never on the outer exposed reef slope. L. kasmira, common
on the outer slope, was never taken in sheltered water. Numbers of species
common to both types of coral area also differed. L. bohar was one of the
dominant species of the outer slope, but although present in the more sheltered
water it was very much less common. Conversely L. gibbus was common and
sometimes abundant in sheltered areas, but on the outer slope it was poorly
represented in the catches.
The reasons for the patterns of distribution described in the above para-
graphs are obviously complex, and not within the scope of this work, but many
of the problems here raised could be approached experimentally. Temperature,
salinity, light, O, concentration and turbidity preferences of the juveniles and
adults of different species could be tested with carefully designed aquarium
equipment, especially for the smaller species, and would undoubtedly give
valuable clues to the reasons for their distribution.
On a typical East African reef the Lutjanids are a major component of
the fish fauna. They and the Serranidae form in general the bulk of the non-
pelagic predators, in contrast to the Carangidae, Scomberomoridae and Sphyraenidae.
They differ from the Serranidae in that while members of this family are usually
often solitary and many are more or less stationary for long periods most
Lutjanids tend to school and move actively over the coral. The commonest
line-caught fish over exposed coral was the large, mainly fish-eating, L. bohar,
with A. virescens common in mid-water and at the surface in the same areas.
The commonest smaller species, L. fulviflamma, L. kasmira and L. lineolatus,
are often present in shoals numbering 50 and more. These species swim close
to the coral and are predominantly crustacean feeders. On sandy bottoms and
Cymodocea beds their place is taken by the Lethrinidae.
In the deeper Lamu Banks off the Kenya coast this family forms the bulk
of the predators, if we may judge by the quantities of line-caught fish. Lutjanids,
mainly L. bohar, L. rivulatus, and L. sanguineus formed 54% of the fishes taken,
the Serranidae 26°% and the Lethrinidae 15%, with 5% of sharks and other
species (Williams, 1958).
No sharply marked breeding seasons were found in any of the species
studied, although sometimes a single sample would contain many ripe fishes
of both sexes. In general all species seemed to breed over a large part of the
year, but mostly in the warm north-east monsoon period.
572 ANNALS OF THE SOUTH AFRICAN MUSEUM
ACKNOWLEDGEMENTS
My thanks for help and encouragement are due to the Director, Dr.
J. F. G. Wheeler and Mr. Frank Williams, Dr. J. F. CG. Morgans and Mr.
B. S. Newell of the East African Marine Fisheries Research Organization,
Dr. P. H. Greenwood of the British Museum, and to Professor J. H. Day,
University of Cape Town, for helpful criticism of the manuscript.
SUMMARY
Seventeen species of the family Lutjanidae from the East African coast
are discussed, and notes on their distribution, feeding, spawning seasons and
shoaling habits are presented.
Eight species of the genus Lutjanus were found only in shallow water, and
were never found below the limit of active coral reef growth (approximately
15 fathoms). The major thermocline at from 25-50 fathoms is suggested to
be a barrier to deeper distribution of the five species (four of the genus Lutjanus
and Aprion virescens) found from shallow water to below the living coral reefs.
Three species (two of the genus Pristipomoides and Aphareus rutilans) were found
only in deeper water, and never above the major thermocline.
No evidence of migrations was found.
Sheltered coral, and coral exposed to violent wave action were found to
differ in the presence or absence of some species of the family, and also in the
numbers of species common to both habitats.
All the species studied were euryphagous predators. Details of feeding are
given.
No sharply defined breeding seasons were found, but the extended periods
in which breeding took place were mostly in the warm months, November to
April.
Regular growth rings were found on the scales of Lutjanus bohar. Checks
were formed at different times of year in different fishes. It is suggested that
these are related to spawning. Growth increments of from 70 mm. (3rd—4th
year) to 35 mm. (1oth—11th year) were estimated.
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ALLEE, W. C., & others. 1949. Principles of animal ecology. Philadelphia: Saunders.
Bowers, A. B. 1954. Breeding and growth of whiting (Gadus merlangus L.) in Isle of Man waters.
JF. Mar. biol. Ass. U.K., 33, 97-122.
CrarKk, J. R. 1958. Consistency of scale reading. Spec. Publ. int. Comm. N.W. Atlantic Fish, 1,
191-192.
EKMAN, S. 1953. <oogeography of the Sea. London: Sidgwick & Jackson.
Fow er, H. W. 1931. Contributions to the biology of the Philippine Archipelago and adjacent
regions. Pt. IV. Bull. U.S. nat. Mus., 100, 11, 1-388.
Harry, R. R. 1953. Ichthyological field data of Rarioa Atoll, Tuamotu Archipelago. Afoll
Res. Bull., 18, 1-190.
Herre, A. W. 1953. Check list of Philippine fishes. Res. Rep. U.S. Fish. Serv., 20, 1-977.
NOTES ON THE BIOLOGY OF THE LUTJANIDAE 573
Hicxuine, C. F. 1933. The natural history of the hake. Part IV. Age determination and growth
rate. Fish. Invest., Lond. (2), 13, 1-120. .
Menon, M. D. 1953. The determination of age and growth of fishes of tropical and sub-tropical
waters. 7. Bombay nat. Hist. Soc., 51, 623-625.
Moreans, J. F. C. 1958. Three confusing species of Serranid fish, one described as new, from
East Africa. Ann. Mag. nat. Hist. (13), 15 643-656.
Moreans, J. F. C. 1959. The North Kenya banks. Nature, Lond., 184, 259-260.
Moreans, J. F. C. In unpublished field data of the East African Marine Fisheries Research
Organization, Zanzibar.
NEWELL, B. S. 1957. A preliminary survey of the hydrography of the British East African
coastal waters. Fish. Publ., Lond., 9, 1-21.
NEWELL, B. S. 1959. The hydrography of the East African coastal waters. Part II. Fish. Publ.,
Lond., 12, 1-18.
RANDALL, J. E. 1955. Fishes of the Gilbert Islands. Atoll. Res. Bull., 47, 1-243.
RANDALL, J. E. 1958. A review of ciguatera, tropical fish poisoning, with a tentative explanation
of its cause. Bull. Mar. Sci. Gulf & Caribbean, 8, 236-267.
RieEpi, R. 1958. An attempt to test the efficiency of ecological field methods and the validity
of their results. In Buzzati-Traverso, A.A., ed. Perspectives in marine biology., 57-65. Berkeley
& Los Angeles: University of California press.
Scuuttz, L. P., & collaborators. 1953. Fishes of the Marshall and Marianas Islands. Bull.
U.S. nat. Mus., 202, 1, 1-685.
SmitH, J. L. B. 1953. Sea fishes of southern Africa. 2nd ed. Cape Town: Central news agency.
SmitH, J. L. B. 1954. Fishes new to Africa obtained by deep line fishing in Kenya waters, with a
revision of the East African species of the genus Pristipomoides Blk. 1852. Ann. Mag. nat. Hist.
(12), 7, 481-492.
STEPHENSON, T. A. 1947. The constitution of the intertidal fauna and flora of South Africa.
Ann. Natal Mus., 11, 207-324.
Steven, G. A. 1930. Bottom fauna and the food of fishes. 7. Mar. biol. Ass. U.K., 16, 677-700.
Sverprup, H. U.; Jonnson, M. W.; & FLeminc, R. H. 1942. The Oceans. Engelwood Cliffs,
N.J.: Prentice-Hall.
Taxsor, F. H. 1957. The fishes of the genus Lutjanus of the East African coast. Ann. Mag. nat.
Hist. (12), 10, 241-258.
Tasor, F. H. 1958. On Plectropomus maculatus (Bloch) and P. marmoratus (n.sp.) from East
Africa (Pisces, Serranidae). Ann. Mag. nat. Hist. (13), 15 748-752.
Tasor, F. H., & NEwett, B. S. 1957. A preliminary note on the breeding and growth of
Tilapia in marine fish ponds on Zanzibar Island. E. Afr. agric. F., 22, 118-121.
Tasot, F. H., & Wiruiams, F. 1956. Sexual colour differences in Caranx ignobilis (Forsk.),
_ Nature, Lond., 178, p. 178.
WHEELER, J. F. G., & Ommanney, F. D. 1953. Report on the Mauritius-Seychelles fisheries
survey, 1948-49. Fish. Publ., Lond., 3, 1-148.
Weser, M., & De Beaurort, L. F. 1936. The fishes of the Indo-Australian Archipelago, 7. Leiden:
Brill.
Wait ey, G. P. 1943. Poisonous and harmful fishes. Bull. Coun. sct. industr. Res. Aust., 159, 1-28.
Wi.uiAMs, F. 1953. Catches of Coryphaena hippurus (L.) in the Western Indian Ocean. Nature,
Lond., 1715 p. 703.
Witutams, F. 1956. Preliminary survey of the pelagic fishes of East Africa. Fish. Publ., Lond.,
8, 1-68.
Wiiuiams, F. 1958a. Fishes of the family Carangidae in British East African waters. Ann. Mag.
nat. Fist. (13), 15 369-430.
WiiuiaMs, F. 1958b. A preliminary report on deep water fishing off the North Kenya coast.
E. Afr. agric. F., 24, 61-63.
Wiurams, F. 1959a. Marlins in British East African waters. Nature, Lond., 183, 762-763.
Wiuiams, F. 1959b. Black marlin in British East African waters. Nature, Lond., 184, B.A.78.
Wiis, F., & NEwELL, B. S. 1957. Notes on the biology of the dorade or dolphin-fish
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ANNALS
OF THE
SOUTH AFRICAN MUSEUM
VOLUME XLV
PART VI, containing :—
A new Solifugid Arachnid from Table Mountain, Cape. Solpuga grindleyi, sp.n.
By A. C. Brown. (With 2 figures in the text.)
SUED JULY 1960 _—~PRICE 1s.
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A NEW SOLIFUGID ARACHNID FROM TABLE MOUNTAIN, CAPE.
SOLPUGA GRINDLEY], SP. N.
By A. GC. BRown
Koology Department, University of Cape Town
Though the Solifugae constitute but a small order of the Arachnida, the
group is extremely well represented in the South African region, 196 species
being now recorded from the area. In particular southern Africa abounds with
species of the family Solpugidae; notably the genera Solpuga and Solpugema,
which account for no less than sixty-five of the South African species. It is no
surprise, therefore, to find that a new species of Solifugid, recently taken on
Table Mountain, in the Cape Peninsula, also belongs to the genus Solpuga.
In contrast to the rest of South Africa, Solifugae are rare in the Cape
Peninsula and only five species, excluding the new one, are recorded from the
area. ‘They are Solpuga fusca Koch (1842), Solpuga monteirot Pocock (1895),
Solpugema vincta (Koch, 1842), Blossiola litoralis (Purcell, 1899) and Toreus
capensis Purcell (1899). The new species, Solpuga grindleyi, is quite easily
distinguishable from each of these, not only in the laboratory but also in the
field. In size and in general appearance it is similar to Solpuga fusca, but whereas
the latter is a black-legged species, in S. grindleyi the legs are coloured yellow-
ochre. The species is very much smaller than Solpuga monieirot and the head-
plate is light yellow-ochre whereas in the latter species the cephalic plate is
very dark and may be almost black. The new species also lacks the median
black abdominal band of Solpugema vincta.
Examination of the chelicerae (mandibles) of both male and female shows
Solpuga grindley to be distinct from any other species of the genus so far recorded
from southern Africa. The dentition of both the upper and lower jaws—not
only the position of the teeth but also their number—at once separates it from
all the other Cape species including S. ferox Pocock (1895), S. schlectert Purcell
(1899) and S. bovicornis Lawrence (1929) as well as the species already mentioned.
The new species closely resembles Solpuga fusca but is distinct from that
species not only in coloration and in the dentition of the mandibles, but also
with regard to the flagellum of the male, that of S. fusca being bzfurcate while in
the new species the flagellar apex is entire.
The author has two specimens of Solpuga grindleyi in his possession. A male
specimen bearing the label ‘Solifugid— Table Mountain’ was discovered among
the class-material used in the Zoology Department for teaching purposes, while
575
S.A.M. VOL. XLV. PT. VI.
CART ISON)
r} AIT HSONIAN Att r g f At
ANNALS OF THE SOUTH AFRICAN MUSEUM
576
Fic. 1
Solpuga Grindleyi, sp. n. Holotype (2) x 10.
A NEW SOLIFUGID ARACHNID FROM TABLE MOUNTAIN, CAPE 577
an adult female was taken complete with nest on the Gamps Bay side of
Table Mountain by Mr. J. Grindley on 20 June 1958. As the female specimen
was studied alive and as the male, though well preserved, does not bear an
adequate label, it has been decided to designate the female as Holotype for
the new species. The male therefore becomes the Allotype.
Solpuga grindleyi sp. n.
DEsCRIPTION OF HOLOTYPE (Female)
Living coloration. Head-plate yellow-ochre with jet-black eyes; chelicerae
light yellow-ochre over most of their surface but giving way to dark brown
towards their tips. Abdomen dull grey at the sides and between segments,
tergites greyish brown; uniformly light grey ventrally. Ventral surface of
thorax light yellow, malleoli yellow with dark brown distal margins. Legs for
the most part yellow-ochre, tending towards pale yellow on the tibiae and
tarsi of the first three pairs. Pedipalps dark yellow-ochre. Setae in general
agreeing with the colour of that part of the body on which they occur, but
setae on the chelicerae dark brown against their light yellow-ochre background.
Apparent proportions. Cephalic plate as broad as long, slightly shorter than
chelicerae. Abdomen approximately four times length of cephalic plate and
twice as broad, evenly rounded. Palp with both tibia and tarsus club-shaped;
setae on pedipalps not arranged in a definite pattern, but several very long
setae on each of the last three segments. First pair of walking-legs slender,
two-thirds length of palp. Second walking-leg somewhat more robust but
shorter than first leg. Third leg stouter than both preceding appendages and
equal in length to first leg. Fourth pair of legs slender except for expanded
femur, longer than all other appendages, including the pedipalp.
Measurements. ‘Total length, 12-4 mm.; greatest (abdominal) width,
3°5 mm. Length of chelicerae, 2:4 mm.; length of head-plate, 2:0 mm.;
greatest width of head-plate, 2.0 mm. Length of pedipalp, 9:2 mm.; femur,
3°0 mm.; tibia, 2-5 mm.; tarsus, 2-6 mm. Length of first walking-leg, 6-6 mm.;
second leg, 5:0 mm.; third leg, 6-8 mm. Length of last leg, 9:8 mm.; femur,
3-0 mm.; tibia, 2-6 mm.; tarsus plus metatarsus, 3-2 mm.
Remarks. The Holotype was taken alive from under a small stone, com-
plete with nest. The latter consisted of a shallow depression lined with soft
chips of wood and a few small pieces of bark. The specimen was kept with
part of its nest in a glass tube for some weeks without feeding. It was then
given a number of small beetle larvae, which it readily devoured. Vision is
apparently poor, as the animal was not disturbed by movements 15 cm. away
from it, except when these movements cast a shadow over it. In the latter case,
and when a finger was moved to within 5 cm. of the animal, it reared up into
578. ANNALS OF THE SOUTH AFRICAN MUSEUM
the defence attitude common among the Solifugae. The entire body is raised
from the ground, the back legs being bent so that the body acquires an angle © :
of some 30° from the horizontal, the pedipalps are elevated and held far
apart, the first pair of legs is lifted from the ground and pointed forwards
and the chelicerae are held agape. The animal orientates itself so as to face
continually the source of potential danger.
THE MALE SPEcIMEN (Allotype)
Colours similar to but somewhat darker than those of the female. Legs and
pedipalps tending towards light brown on tibiae and tarsi of all appendages |
Fic. 2
Solpuga Grindleyt sp. n. Male Chelicera x 40.
and also on the femur of the hind-most leg, though darkish yellow-ochre still
predominates. Proportions of parts as for female except that the abdomen is
slightly narrower; total body length, 13:6 mm. Body and appendages more
setose than in Holotype and the setae themselves very much darker, varying
from light brown on the legs to very dark brown on the head-plate and
abdomen.
Male chelicera as shown in figure 2. Flagellum very long and slender,
being approximately twice the length of the chelicera itself; flagellar shaft
tapering evenly, without spicules or denticles. Ventral jaw with a large blunt
tooth proximally, followed by two less prominent rounded processes; a convex
surface leads to the curved and tapered fang. Dorsal jaw with two large rounded
teeth preceding the fang-tip. Between the proximal tooth and the articulation
A NEW SOLIFUGID ARACHNID FROM TABLE MOUNTAIN, CAPE 579
of the lower jaw occurs, after a short adentate region, a double row of proceses
- between which the lower jaw fits when at rest. The inner row of processes
consists of a flat-topped projection distally to which are attached four stout
elongate plumose setae pointing forwards, followed by three sharp spines. The
outer row projects somewhat lower than the inner row and consists of two
naked flat-topped processes, the first (distal) process being much larger than
_ the second, followed by three sharp spines. Five extremely long, stout, plumose
setae are attached immediately behind the base of the flagellum and lie over it,
pointing forwards. The distal halves of both jaws are without setae but the
proximal part of the lower jaw is quite heavily setose, the setae on the ventral
side being simple, those towards the upper surface plumose.
Both types have been deposited in the South African Museum.
REFERENCES
Koch, C. L., 1842. Die Arachniden. Arch. f. Naturg., 8 (1).
Lawrence, R. F., 1929. New South African Solifugae. Ann. S. Afr. Mus., 29 (1), pp. 153-179.
Lawrence, R. F., 1955. Solifugae, Scorpions and Pedipalpi in South African Animal Life, 1,
pp. 152-262 (Uppsala). :
Pocock, R. I., 1895. Notes on some of the Solifugae contained in the Collection of the British
Museum, with Descriptions of New Species. Ann. Mag. nat. Hist. (6), 16, pp. 74-97.
Pocock, R. I., 1897. On the Genera and Species of Tropical African Arachnida of the order
Solifugae, with Notes upon the Taxonomy and Habits of the Group. Ann. Mag. nat. Hist
(6), 20, pp. 249-272.
Purcell, W. F., 1899. New and little-known South African Solifugae in the collection of the
South African Museum. Ann. S. Afr. Mus., 1 (3), pp. 381-432.
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The ANNALS OF THE SOUTH AFRICAN MUSEUM are issued in parts at irom 4
-
intervals as material becomes available.
Out of print:
VI (Part 1, Index), VII (Parts 1-3), VIII, IX (Parts 1, 2),
Vol, 1, “UL: (Parts) 1-955; 92\0), Leb (Part 1) x7
XXI (Part 1), XXIV (Part 2), XXXI (Part 1).
Current prices:
Vol. II.
Dit
IV.
XXII.
yo, Ole
XXIII.
XXIV.
XXV.
XXVI.
XXVII.
XXVIII.
XXIX.
XXX.
INDEX
XXXI.
XXXII.
XXXII.
XXXIV.
XXXV.
XXXVI.
XXXVITI.
XXXVIII.
XXXIX.
ole
XLI.
XLII.
XLIII.
XLIV.
XLV.
1900-1902
1903-1905
1903-1908
1906-1910
1908-1910
1908-1913
IQiI-1918
IQII—1914
IQII—-1918
IQI3—-1924
1913-1923
1915-1924
1914-1916
1917-1933
1917-1920
1921
1924-1925
1924-1926
1925-1927
1925-1928
1925-1926
1929-1938
1927-1928
1928
1929
1929-1932
1929-1931
1931-1935
Zoology and Geology (excl. Parts 1-3, 5, 7, 8)
Zoology (excl. Part I)
Palaeontology
Geology, Palaeontology, Zoology, Anthropology (excl.
Parts 4,2,)5, 05.0)
Zoology (eter Part I, Index)
Palaeontology (excl. Parts 1-3)
Botany (excl. Parts 1, 2)
Zoology (excl. Part 5)
Zoology (excl. Parts 2, 7, indeon
Palaeontology and Geology
Archaeology and Zoology
Zoology ss oe
Zoology
Botany
Zoology
Zoology
Zoology
Zoology
Zoology eet Part o
Palaeontology
Zoology ;
Anthropology and Fitnoto ey geet Part be
Zoology : if = ig
Zoology
Anthropology
Palaeontology
Zoology
Zoology
of papers, authors and subjects pnblichea in Wolls. I- XXX
1934-1950
1935-1940
1939
1938
1956
1942-1948
1947-1952
1950
1952
1952-1956
1952-1955
1953-1956
1955-1957
1957-1959
1959-
The LIBRARIAN, Soutn Arrican Museum, Care Town.
Palaeontology (excl. Part 1)
Zoology
Zoology
Zoology
Zoology
Zoology ;
Agchatoloiny
Zoology
Zoology
Botany
Zoology
Palaeontology
Zoology and Palacomiglony.
Zoology and Palaeontology
Zoology and Palaeontology Part 1
Part 2
Part 3
Part 4
Part 5
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