37 

Palaeont. afr., 15,37- 64. 1973 

A REVIEW OF THE EVIDENCE FOR MARINE CONDITIONS IN 
SOUTHERN AFRICA DURING DWYKA TIMES 

by 

I. R. McLachlan and Ann Anderson 

Bernard Price Institute for Palaeontological Research 

Abstract 
General Geology 

Kalahari Basin 
Warmbad Basin 

CONTENTS 

Great Karroo Basin .... 
Marine Incursion into South West Africa . . . . . 
Previous Fossil Evidence for a Marine Incursion into the Great Karroo Basin 
A Marine Dwyka Fauna from the Great Karroo Basin 

Location 
Stratigraphy . 
Description of the Fossils . . . . . . 

Reconstruction of the Palaeo-environment During Dwyka Times 
Tillite Stage . . 
Marine Horizon at the Base of the Upper Dwyka Shales Stage 
White Band at the Top of the Upper Dwyka Shales Stage 

Conclusions 
Acknowledgemen ts 
References 

ABSTRACT 

37 
37 
37 
40 
40 
40 
40 
41 
41 
41 
45 
45 
45 
46 
46 
47 
47 
47 

The faunas of the three major Dwyka basins in southern Africa are listed in the form 
of tables. Marine invertebrates have been recorded from the western part of the Kalahari 
basin and from the Warmbad basin, but had not previously been confirmed within the 
Great Karroo oasin. A new fauna from the base of the Upper Dwyka shales near 
Kimberley is described. Cephalopods, brachiopods and lamellibranchs are found in 
calcareous concretions, which also contain palaeoniscoid fish, coprolites, fossil wood and 
the mineral glauberite. The marine invasion into South West Africa postulated by Martin 
& Wilczewski (1970) therefore extended into the Great Karroo basin as well. By the 
time of accumulation of the White Band at the top of the Upper Dwyka shales, 
conditions were probably non-marine; with the possible exception of the Notocarid 
crustaceans, the White Band fossils are not, in themselves, indicative of marine conditions. 
The only other significant indication of marine conditions in the Great Karroo basin is 
the glauconite in the deltaic Coal Measures of the Ecca in the northern part of the basin. 
It is possible then that the fossiliferous marine shales near Kimberley accumulated as a 
fine-grained offshore facies of the Ecca deltaic sequence. 

GENERAL GEOLOGY area and eastern Botswana, the central portion of 
the basin being covered by Cenozoic Kalahari 
sediments. Green (1969, fig. I) illustrated the 
continuity of the Karroo strata between the 
eastern and-western outcrops. The Dwyka series is 
well exposed in South West Africa, but in eastern 
Botswana it is poorly exposed and is of variable 
thickness and discontinuous development. A 
white-weathering unit at the top of the Dwyka 
sequence contains Mesosaurus, and is considered to 
be the equivalent of the White Band (du Toit, 
1954, p. 329; Martin, 1961a, p. 34; Heath, 1966, 
p.42)(see also fig. 1 and table 5). Coals occur 
below this horizon in the Upper Dwyka Stage 
(Martin, 1953; 1961a, p. 34). 

The extent of the Dwyka sediments con­
sidered in this article is shown in fig. 1. Generalised 
stratigraphic sections of the three main basins are 
given in fig. 2. For the purpose of this discussion, 
the White Band is taken to mark the top of the 
Upper Dwyka Stage. 

Kalahari Basin 
(Name from Martin, 1961a, p.15. Synonyms 

include: "South Kalahari basin", Martin, 1961a, 
table II, 1968, table 2, 1970, p. 225; "Kalahari 
Karroo basin", Green, 1969, fig. I; "Botswana 
block", Frakes & Crowell, 1970, p. 2263.) There 
are outcrops in South West Africa, the Gordonia 



24' 

26' 

28' 

30' 

South 

West 

Africa 

,.;,.;- - ------

KALAHARI 

22' 

" --,~' 

BASIN 

i 

" , , 

(/ .--------------------------- ----~' 
I \ ~~tswana //,~ 
\ -~ 

I 
I 

'. 

.... 

~ ------- -- " ,<~:>~-~--::---.>' ,'~ C:i ~ ___ .. __ --._:..... ,"" ','iJ" ,W ---.,. "-, '" " " IA \ '. 
.. ~ •• I •• , .. ,., -', -', " " ',"",," ,. 

.'. " ' 0- -'- " " " '. .. '\ Q 

• • ..... . ' ' _" 0 
, " ....... ......... , ... , \g I 

.,.'.. a... ' " • '.'''' ' 
, ....... ~ " " "''' , . ' • .' .' ••• _ • ..... '<b '" ... ,\'6 " , ' '. , ... ~ '. ~ 0-" ." , " -'./ , ,,">"- • ~,' .' " ,._-100---- -' 
.. \ ....... '. """" --- ' ' ' _, ...... \'i."',,"R.DE U .,,' ", ' ' / 
, 0(1';t;L. ~ "',., KAR ' \AI,' I I I. I J , /---:h._ / / , / ,~ • , ROO" " " ,'- ,,/ 

". ...., ',', " ! ASI.!::: / ... -- " / / '.0._" • ." · " " ' ' , ,," -' ,---.... ,!' 
__ ._ ' .... " " " ' ........ , ___ " ;' ,',' I" .,' vv.., f I ;' !b!!! ,.t........ __ - "I'" ... ' I . . .. .,,' , , ,,_ " , !, __ ..... ' I' , ,. .... , . ' ',' -",' ' -- ' ' " ' . . . . .,,' ,,'~ .. ---- " " ': ' -- . " ..,' , _ _ -'.to- ,......... ___ ........, I " I I I / /. VR 1 \ J" I 

... '''' " ' ,,' ' " ., ".' ", """, "" , . ,- - ',' '--' , ' , ' , ' " ' 
__ .. __ _, l"'" " ",' , "" " , 
--::C:'C> • ........... ,,,-,,=,-' J ,'," I'" ' ' ._ "" ,_' C ' . ·r- ',-/ -', ',' ' ,/" ' , 
_--- _- ,,_<F0 / / ,- - -- --" • ,', / / ' " " ,.....- -- .. ' / " , .. ' , / ' ' , . 
"..- /',,0" ,/ " , ' ' , ,:.,.. ' ' '.' 

Soutn Africa 

22' 24' 26' 



• Durban 

... 100--. 

e!1u 

® 
® 

© 

Fig. I 

EVIDENCE FOR A MARINE I NCURSION INTO 
SOUTHERN AFRICA DURING DWYKA TIMES 

o 50 '00 '50 miles 
or. --5',...0---J· ........ 'o,...0--, .... s .... d--2"Tcl-o-..... ;so km. 

International boundary 
Town 

Extent of Dwyka (after Haughton, 1969) 

Elevated areas (after Stratten, 1968) 

Upper Dwyka Shale isopachs in feet (after Winter & Venter, 
1970) 

Marine fossil site -AB 1 Borehole yielding Acritarchs 

? Radiolaria (after !ltrydom, 1946) 

3 

Appoximate locality of Mesosaurus spp. (after Haughton & Frommurz 
1927, Martin, 1953 & 1961 and from the South . Afritan Museum, 
Cape Town ((S.A.M.)) and Alexander McGregor Memorial Museum, 
Kimberley ((A.M.M.M.l) collections) 3 

Approximate locality of crustaceans Ntt~cgriS tapscotti 
Pyggspis ginsburgi (S.A.M. and A.M.M. . 

32° 34° 36° 

or 



40 

Warmbad Basin 
(Name from Martin, 1961a, p. 34. Synonym: 

"Orange River Basin", Frakes & Crowell, 1970, 
p. 2268.) This outcrop is now quite distinct, but 
Martin & Wilczewski (1970, p.227) suggested 
that it might have been connected to the north 
with the Kalahari basin before the rise of the 
Karasberg horst mountains. There may also have 
been a connection with the Kalahari basin outcrop 
in Gordonia, as the present break is probably due 
solely to erosion by the Ham river. The Dwyka 
series is well exposed in the Warmbad basin, and 
includes a characteristic White Band at the top of 
the sequence. 

Great Karroo Basin 
(Name from Martin, 1961a, p. 30.) The basin is 

separated from the Kalahari basin along nearly the 
whole of its length by the Transvaal highlands and 
Kaap plateau. There was apparently a gap between 
the Kaap plateau and the hypothetical "western 
highlands" through which the Great Karroo basin 
extended (see fig. 1); at present it is only separated 
from the Warmbad and Kalahari basins by the 
erosional valley of the Orange River. Dwyka tillite 
is ubiquitous in the southern part of the Great 
Karroo basin, and is also preserved in the deeper 
basement valleys in the north. 

Much of the "tillite" in the northern part of 
the basin is possibly glacial material reworked 
during later sedimentary cycles (du Toit, 1921, 
p. 217; 1954, p. 274; Stratten, 1968, p. 26). The 
White Band of the southern Karroo is recognised as 
far north as Hopetown in the west and Port St. 
Johns in the east (Stratten, 1968, p. 20), but it is 
probable that it does extend beyond these points. 
Mesosaurus spp. and Notocarid crustaceans, 
typical of the White Band in the western Cape, are 
found in White Band-like shale farther north; e.g. 
Mesosaurus sp. (A.M.M.M. 4431) collected from 
Kameelpan, 32 km north-east of Kimberley, and 
Notocarididae (A.M.M.M. 2689, 3689) from the de 
Beer's mine property in Kimberley. No coal is 
known to occur below the White Band, and this 
horizon nowhere directly underlies the Middle 
Ecca Coal Measures. 

MARINE INCURSION INTO 
SOUTH WEST AFRICA 

In this article the word "marine" is used in the 
biological sense; we do not consider a large body of 
water necessarily to have been marine, unless there 
is supporting palaeontological evidence. 

Marine invertebrates have been known from 
the Kalahari basin in South West Africa since the 
beginning of the century (Schroeder, 1908; Range, 
1912; du Toit, 1915; Dickins, 1961; Heath, 1966; 
Martin, Walliser & Wilczewski, 1970; Martin & 
Wilczewski, 1970; Wass, 1972.)(see table 1). 
Dickins (op cit.) described collections of Eury-

desma my tZ"loZ"des and Peruvz"spz"ra vZ"persdorfensis, 
and concluded that they were of Lower Permian 
age, most likely Sakmarian, but possibly early 
Artinskian. 

In 1927, Haughton & Frommurze reported a 
marine lamellibranch (described by Reed, 1935) 
and a very poorly preserved gastropod from Haib, 
in the Warmbad basin, while Martin & Wilczewski 
(1970) recently recorded foraminifera from 
Kanibeam. Our discoveries of palaeoniscoid fish, 
lamellibranchs, radiolaria, foraminifera and sponge 
spicules at Klipneus and Zwartbas have extended 
the fauna from that basin (table 2). These fossils 
are confined to the lower part of the Upper Dwyka 
shales (see fig. 2). 

The arthropod walking trails first reported by 
Haughton & Frommurze (1927) from flagstones, 
including varved units, in the lower part of the 
Warmbad basin section (immediately above the 
tillites, but below the marine faunas) are similar to 
the assemblage from varved shales within the tillite 
in Natal (cf. Savage, 1970, 1971). These rich trace 
fossil faunas are not, in themselves, indicative of 
either marine or fresh water conditions. Indeed, 
the varves imply deposition in non-saline water 
(Duff et al., 1967, p. 50). 

Martin & Wilczewski (1970, fig. 4) proposed 
a marine embayment from the west, open to the 
ocean from the Windhoek highlands south to 
include at least the Warmbad basin, and extending 
for an unknown distance east into the Kalahari 
basin. They did not extend the embayment into 
the Great Karroo basin because of the lack of 
palaeontological support. 

PREVIOUS FOSSIL EVIDENCE FOR A 
MARINE INCURSION INTO THE 

GREAT KARROO BASIN 

There have been various speculations on the 
extent of the sea that supported the South West 
African marine faunas. Haughton (1919, p. 10) 
suggested that . ". .. the presence of marine 
mollusca in the Upper Dwyka of the South-West 
Protectorate leads to the supposition that the 
Upper Dwyka clastics of the Union [of South 
Africa] were probably laid down in an estuarine 
arm of the sea". Du Toit (1921) and Stratten 
(1968) enlarged on this concept. 

The few known records of animal fossils from 
the Dwyka sediments in the Great Karroo basin are 
summarised in tables 3, 4 and 5. Apart from the 
White Band faunas, the only invertebrates previous­
ly reported from the Dwyka are "radiolaria" from 
the Laingsburg area. These reports were tentative 
and we do not feel that the presence of radiolaria 
has been demonstrated. For a more detailed 
discussion see table 3. 

The White Band, at the top of the Upper 
Dwyka Shales, has yielded the fish Palaeonz"scus 
capensis (table 4), Notocarid crustaceans 
(Notocaris tapscotti and Pygaspis ginsburgi [table 



41 

KA LAHARI BASIN (Mariental-A. ab) 
(after Heath.1966. See fig . 3 ) 

WARMBAD BASIN GREAT KARROO BASIN (S.cape) 

(a ft er Winter& Venter. 1970) (a fter Haughton & Frommurze,1927) 

_ Me. o50uru. 

; ... '" Cl 0 -
-,):-<t c 0- 0 0 VI 

<t ;:c( 
:..: .;~ >- -VI 
~ c -
O 0 

U 
<C ... 
"-
"-
::> 

~. - -o -c -- -e __ 
c _ 
.~ - --
0 - -
\of _-_ 

~ 
-

c ---
~ -
~ - -

e - -
c -
0 _-_ 

White Band 

i----- Me.o.aurus _ _____ ., 

Crustaceans 
~. - -
o c - _ -e __ 
.~ -

Carbonaceous shale w ith thin ~ = -:::: 
impure limestone bands and phosphatic .. - = -
lenses and cone, . tions 

-- -- ~ ----- - .~ -=--
. :: Nossob Sandst~ ___ _ - -

------7t_..,..~=~= ' unconformity L 
-_ A - (SW.A. only dashed line indicates 

... 
Cl 
~ 
VI 

.. 
. ~~ 
o c _0 

~- - top of glacial beds 
04. _ 0 

_ 0_ 
o _ 

A 
o . -

Max . total 
thickness SOOm 

( Mortin. 1961 ( b~ p 200) 

Tillite and boulder mudston-;- - _~ - _ -

with some Interbedded 
carbonaceous shale and 

varved shale 
A 

A 

A 

A 

A 

'" 

A 

A 

A 

A 

o 

so 

100 

200 

300 

GENERALISED STRATIGRAPHIC SECTIONS 400 

Fig. 2 

3 and fig. 1]) and the small ' aquatic reptile 
Mesosaurus (table 5 and fig. 1). From our 
observations in the Loeriesfontein area, it seems 
that the fish occur in finely bedded shales near the 
base of the White Band, while the crustaceans 
occur in poorly bedded sediment near the top and 
Mesosaurus in a silty, bedded facies , also near the 
top of the unit. Mesosaurus has been regarded as 
both a non-marine (Holmes, 1965, p. 1222; 
Romer, 1966, p. 117) and a marine animal 
(Rayner, 1970, p.475; Teichert, 1970, p.132; 
Meyerhof & Meyerhof, 1972, p. 290). In isola­
tion, the skeletal remains do not proclaim the 
salinity of the water inhabited. However, from the 
available associated evidence, there is little to 
suggest that the water was marine (see below). 

Clear fossil evidence of marine conditions In 

the Great Karroo Basin was therefore lacking. 

A MARINE DWYKA FAUNA FROM 
THE GREAT KARROO BASIN 

There are undescribed specimens of the 
palaeoniscoid fish? Namaichthys schroederi in the 
Alexander McGregor Memorial Museum 
(A.M.M.M.), Kimberley. The fish were collected by 
Robert Broom, in about 1940, from the farm Zand 

metres 

Bult near Douglas. They occur in concretions very 
similar to those containing marine invertebrates in 
South West Africa. We revisited Broom's site, and 
found that the concretions also contain marine 
invertebrates. More recently we discovered a 
second site near the Tankwa river (fig. 1) at the 
same stratigraphic position as the Zand Bult 
(Blaauw Krantz) locality. No invertebrates have yet 
been recognised, but wood, fish and coprolites are 
present. Only the Blaauw Krantz site is described 
in detail here. 

Location 
The richest fossil site has been named after the 

farm Blaauw Krantz. A single fragmentary 
cephalopod was found on the farm Zand Bult, and 
several large fossilised tree trunks occur in the 
shales on the farm Olie Rivier (fig. 4). These 
properties are subdivisions of the original farm 
Zand Bult. 

Stratigraphy 
Du Toit (1915, pp. 44-45) noted that "The 

Dwyka in South West Africa is more or less a 
replica of that obtaining along the strip of country 
extending from Vryburg to Prieska, and the 
conditions which prevailed during its deposition in 
the former region appear to have extended up this 



42 

..: ... 
0 ... 
... Z 
;:)0 
0 ... 
oj! 11'1 

o 
11'1;:) 

~~ ... 

DWYKA SECTION IN THE MARIENTAL-ASAB AREA, S~~. 
BASED ON HEATH 1966. 

UNIT NAMES AFTER HEATH , 

White Band? 

Upper Shale 

Auob sandstone 

Grey shale 

Nossob sandstone 

Upper boulder 
mudstone 

~ 50ft Mesosaurus & fossil wood 

~500' 

Fossil wood .. '. 
+ 140 ~ ~ > ~ Thin coal seams - :-:-:;:.:0 0 . ,0: .. ' 

~200' 

t SO' ::::: : : : ::: ___ Fossil wood 

--- - - - _Gasteropod sp~Maack,pers.comm.toMartin) 
t 110' - - - - ~ ---1Qrthoceras sp.(du Toit)Ts., 

- 0...... - Iblryd.sma sp(Schroed.r),ltzawisis 
~----------------;-----~~ 

... 

... 
z ... 
~ 
Z 
0 
..: 
> 
Z ... 
..: ... ... « 
~ 
x 
!!! 
:.t 
U « ..: 
oj! 

0 ... 
x 
11'1 ... 
..: ... 

Lower boulder 
mudstone 

-::.. -= i Euryd.sma slobosum(Range),Gaus 
+-150' -_ -_A -_ Gasteropod sp(Martin1953) Betwe.n AsabtTses 

Crinoid indet. - ..'.. ..-
- _0-_ -_ :- Peruvispira vipersdorfensis(Dickins),Viperstorf 

I--------------+------t Forqminif.ra~scol.codonta,sponge & .chinoid 
- - - spln.s(Marhn&Wikz.wski,1970) 

Yellow green 
mudstone 

- - -- - Eurydesma mytiloid.s(Diclcinsl 
~26ci - _---Brachiopod indet.(H.athl - --V Bryozoan ind.t. .. = ::: Asteroidean ind.t. .. 

Hardap 

;. ..... :-;-=:.:::: . ..... Pala.oniscoid fish(GOrich), Ganikobis 
h: Z .... 40-n-.-o .... f-a .... llt-.-rn-a....,.t:-in-g--+-S-0 ...... ,........"" .. -,-~-----41~ ~ Con~lari~ ind.t.(Schroed.r)," . 
sandston.&mudstone- ..... ' Radiolaria (M&W1970~ - ,Schllp, 
F-==i=F=:::::.:;';;';~--li·=·~·~·~ Goniatit. .... -
Zone of alt.rnahr.g ""=~8 0 o~ Wood .... _ 
tilloid & small ~ 120 0 S) 
boulder mudstone . .? . 
L II b Id - -- - ~ GlaCiomarine. and ground moraine. 
ow.r sma ou er +SO No fossils 

mudstone - A A 

Basal tillite 

Basement 

A 
A A 

t120A A A 

WW/ 
N-I/'//////.-S 

1&.1 

" c( 
I-
In 

c( 
~ 
>-
~ 
0 

cae: 
1&.1 
a. 
a. 
;:) 

Fig. 3. It is not clear how the member names used by Martin & Wilczewski 1970 apply to the units 
mapped by Martin 1953, or Heath 1966. Martin (written comm. 1973) says that the three 
glaciomarine units thin towards the south until the Ganikobis shale comes to rest on the basal 
tillite. 

pre-Karroo depression to which I have gIVen the 
name of the Kaap Valley." 

The basement in the area is formed of 
Precambrian Ventersdorp lava that possessed a 
moderate relief at the time of Dwyka accumula­
tion. As the basement does not outcrop at any of 
the sections measured, the total thickness of the 
Dwyka tillite could not be determined. It is likely 
to vary considerably due to the uneven nature of 
the basement. The top of the Dwyka tillite has 
been put at the level of the highest dropstone­
bearing bed, giving an approximate thickness of 

Fresh exposures near Douglas are largely 
confined to the banks of incised rivers. A 
composite section (fig. 5) has been compiled from 
four locations (see fig. 4) near the Blaauw Krantz 
fossil site. Unfortunately, the strata between the 
sections are obscured by recent sediments, and 
their relative vertical placings had to be made on 
lithological similarity and projected geology. 



43 

LOCATION OF FOSSIL SITES NEAR DOUGLAS. 

Scale 1: 250,000 

°t-I _ .... 1 r-..... ~ ......... ,--"-~ ---.,.J'--ii miles 
o ':1 4 ;, 8 km 

N 

t 

---

". Ki",ber\~Y::,=;­
·w-SO""' 

i' 
\ --"::..­

Ki",b.!! ~- ---

E XPLANA liON 

- -"' 
Railway 

Road 

Farm boundary 

Outcrop 01 los sililerau. concretion 
bearing shale examined 

Fig. 4 

37,5 m for this interval. The predominant rock 
type is dark grey shale, often varved and containing 
dropstones in varying concentrations. Striated and 
faceted pebbles or boulders are usually present. At 
De Kalk this bedded diamictite ("Shaley tillite"; 
Stratten, 1968, p. 33) is contorted, most probably 
by slumping prior to consolidation. Impersistent 
iron and carbonate-rich shale bands, up to 60 em 
thick, are common; they may pinch out laterally or 
grade into bright yellow-weathering cone-in-cone 
limestone. 

The total thickness of the Upper Dwyka 
Shales has not been determined as the White Band 
does not outcrop near the fossil sites. The section 
is directly overlain by a variable thickness of 
calcrete-cemented Quaternary sediments, topped 
by red Kalahari sand (figs. 5, 9). At Blaauw Krantz, 
approximately 18 m of dark grey splintery shale is 
exposed. A dolerite sill intrudes the upper portion 
of the shale. A 6 m zone containing calcareous 
fossiliferous concretions lies 9 m above the base of 
the shales. The concretions occur in bands at four 
levels. They protrude from the softer shale, and are 
concentrated in small gullies by erosion. Their 
shape is commonly oval and flattened, but in the 
lowermost band they are frequently irregularly 
lobed. When fossils are present, they usually lie on 

FOSSil SITES 

1 Blaauw Krantl 
2 Zand Bult 
3 Olia Rivier 

STRATIGRAPHIC SECTIONS 

1 Blaauw Krantl 
"De Kalk 
5 Riet River Bridge A 
6 Riet River Bridge B 

a bedding plane near the middle of the concretion, 
and may extend through the side of it. Weeks 
(1935) provided a convincing explanation of the 
formation of such concretions containing 
uncrushed fossils. He argued (op. cit. p. 171) that, 
in an environment generally unfavourable for pure 
limestone deposition, calcium carbonate concre,­
tions can develop locally around, say, an ammonite 
or a fish, because where there is anaerobic 
decomposition there is a localized concentration of 
ammonia or amines. This would markedly increase 
the pH, sufficient to precipitate the carbonate. The 
original shell or bone material of the fossils from 
Blaauw Krantz is sometimes IlTeserved but, more 
usually, it has been remove-d by solution and 
replaced by crumbly yellow limonite. 

An interesting feature is the occurrence of 
hollow moulds of crystals (approximately 10 em 
long and 2-3 em wide) in some of the concretions. 
The crystal casts were identified by Loock as being 
of the monoclinic salt glauberite (Na2 S04' 
CaS04 ). Glendonites, which are calcitic 
pseudomorphs after glauberite, have been de­
scribed from very much the same stratigraphic 
position in Permo-Carboniferous sediments in New 
South Wales (David et al., 1905), Queensland and 
Tasmania (Raggatt, 1937). 



44 

COMPOSITE STRATIGRAPHIC SECTION 
Vertical scale 1cm. 3metres 

RECENT TO 

QUATERNARY 

I 

BlAAUW KRANTZ 

1 

Weathered grey shale 

1-----
~~~ 

0000 

UPPER DWYKA Ib ~o~ 
SHALES 

] 

Fossiliferous concretion zone 

in weathered grey shale 

489 

~==== e 486 

___ .487 

488 

T~ JU. v!uy~ ? ..ilia.l!... _ 
with • drops tones 

" Very laint indeterminate 
invertebrate trails 

RiEl RIVER 8RIDGE 
6 

xx x-

6 

~ 

I 
e 487 

A 

EXPLANATION 

Refers to location of section on Fig.4 
Kaiohari sand 
Calcrete 

Doh"ite 
Dark friable shale 
Hard iron-rich calcareous band 
Fossiliferous concretions 
Yellow -weathering cone -in-cone limestone 
Varved shale 
Dropstones in varv.d shal.s 

Palynology sample number 

RiEl RIVER 8RIDGE 

5 

8 

--= 
This band 

TriicedTateraily 

DWYKA TILLITE 

These bands converge 
and join 200tt away 

DE KALK 

4 
r;.~..:...= 

_ - .o.~ ' ~' 

----I.~~~ 
--?- - - -~~=-:: 

1--
483. 1----

:= ---=: 
1- =-
1- =--
1--_-
I-- -

Hard unfossiliferou. ~ ~­
carbonate nodules I E __ ~_ 

1- -
4114 1- ---

48 

Folds of amplitude 10 It 
caused by slumping 

IV\.I'Y\ 



Description of the Fossils 
The number of specimens collected of each of 

the fossil groups (table 3) does not reflect their 
true relative abundance, as some survive weathering 
better than others. Brief descriptions of the various 
fossil groups follow: 

(i) Cephalopods (figs. 12- 16) 
One specimen came from Zand Buit, and eight 

from Blaauw Krantz, where most are from the 
lowermost concretion band. The specimens consist 
of portions of the external moulds of 
phragmacones, which show a fine ornamentation 
oflongi tudinal and annular striae (see fig. 15) . Two 
specimens are of internal and external moulds of 
the body chamber with portion of the phragma· 
cone (see figs . 12-14). The body chamb er is 
strongly flattened and the siphuncle is centrally 
placed. The original nacreous shell material is 
occasionally preserved. Teichert confirmed, from 
stereopair photographs, that they belong to the 
order Orthocerida. 

Du Toit (1915, p. 45 ) found a fragment of a 
nautiloid cephalop od Orthoceras sp. near Tses in 
South West Africa (see table 1) . The specimen is 
insufficiently well preserved to warran t close 
comparison with the Blaauw Krantz material. 

(ii) Lamellibranchs (figs. 22- 26) 
Only two of the several hundred concretions 

examined at Blaauw Krantz contained clearly 
recognisable lamellibranchs. Dickins has kindly 
identified a nuculanid, Phestia sp. (figs. 24-26) 
and, tentatively, a nuculid, Nuculopsis sp. (figs. 22, 
23 ) from stereopair photographs. The Phestia sp. 
apparently belongs to the P. darwini group of 
species "... which are fairly widespread in the 
Lower Permian, although similar shells also appear 
to be present in the uppermost Carboniferous." 
(Dickins, 1972, written comm.) 
(iii) Brachiopods 

Only one concretion, (1.40), yielded 
brachiopods. They occur as external casts. Dickins, 
using the specimens and latex casts, has identified 
them as Attenuatella sp. The genus is apparently 
confined to the Permian, and occurs in New 
Zealand, Australia, North America and the 
U.S.S.R. 

(iv) Fish (fig. 27) 
Fish remains are relatively common. Scales 

and the larger skeletal elements, especially the 
skull, are often well preserved and undistorted. 
Jubb has identified the Kimberley museum 
specimens as most probably being Namaichthys 
schroederi. This species occurs in similar deposits 
in the Warmbad and Kalahari basins (table 4). It 
appears from the association of this species with 
marine invertebrates at the same horizon at Blaauw 
Krantz that the fish lived in a marine environment 
for at least part of its life. This is supported by the 
presence in two of the concretions (nos. P2 and 
P6) from Zwartbas, in the Warmbad basin, of N. 
schroederi with arenaceous foraminifera exposed in 
the matrix (see table 2). 

45 

(v) "Spiral coprolites" (figs. 18- 21) 
Together with the fish, these are the most 

abundant fossils at Blaauw Krantz. They are 
generally well preserved, and many of the speci­
mens are complete. Less well preserved examples 
occur in the same stratigraphic position at Klir.neus 
and Zwartbas in the Warmbad basin (table 4 . Our 
tentative identification of the specimens as spiral 
coprolites has been confirmed by Williams. He 
regards the form shown in fig. 18 (1.20) as being an 
"enterospiron" or the fossilised contents of an 
intestinal spiral valve, probably of a shark 
(Williams, 1973, written comm.). This is the first 
indication of the class Chondrichthyes in the 
Permian of southern Africa. 
(vi ) Fossil wood (figs. 10, 11) 

Fossil wood is common in all the concretion 
bands at Blaauw Krantz. Several large tree trunks, 
up to 4 m in length and 25 cm in diameter, are 
exposed at Olie Rivier. The ends of most of the 
pieces of wood are noticeably rounded (fig. 11) . 
(vii ) Palynology 

Samples were processed from the positions 
indicated on fig. 5. Miospore yields and preserva­
tion were generally poor due to the adjacent 
dolerite, but the assemblages were compatible with 
the Lower Permian (Sakmarian) age suggested by 
the molluscs Q. M. Anderson, pers. comm.). No 
acritarchs were n oted. 

RECONSTRUCTION OF THE PALAEO­
ENVIRONMENT DURING DWYKA TIMES 

Tillite Stage 
Stratten (1968) considered that the tillite in 

the south represents a ground moraine of terres trial 
origin. In the north-west, from Loeriesfontein to 
Vryburg, the glacial sediments are bedded, indicat­
ing subaqueous deposition, but whether or not the 
water body was marine has not yet been 
established. Intercalated varves occur within the 
tillite in the southern Karroo (Stratten, 1968, 
p. 111), at the Douglas fossil sites and also in valleys 
in the northern Karroo basin (Cousins, 1950, 
p. 233; Hunter, 1969; Savage, 1971, p. 217). These 
were probably accumulated in fresh water; the 
development of varves is apparently inhibited by 
saline conditions, due to flocculation of the clay 
particles (cf. Pettijohn, 1957, p. 163; Duff et al., 
1967, p. 50)(HaIbich, 1958, pp. 119- 121; 
Rowsell, 1969, pp. 77-81 notwithstanding). 
Acanthomorphitae acritarchs (15-30 microns in 
diameter) have been recovered from shale intercala­
tions in the lower 200 m of tillite in the southern 
Karroo boreholes (OL, SA, VR) shown in fig. 1 Q. 
M. Anderson, in prep.). Without supporting 
evidence, they should not be taken to indicate 
marine conditions. Shaley bands at several 
positions within the tillite stage in the southern 
Karroo and in pre-Dwyka valleys farther north 
(borehole U.C. 608)0. M. Anderson, pers. comm.) 
and near Douglas (see fig. 5) have yielded 



46 

moderately abundant miospores and coarse 
cuticular material. This suggests that vegetation 
was established within the Karroo basin during the 
period of glaciation, and supports the evidence, 
summarised by Plumstead (1969, p. 34), of leaf 
and stem remains within the glacial sediments. 

The Kaap valley (Vryburg-Douglas, see p. 5) 
was probably occupied by a large glacier which 
originated on the Transvaal highiands and the Kaap 
Plateau. The western side of the valley rises sharply 
to a prominent scarp. 

Marine Horizon at the Base of the Upper Dwyka 
Shales Stage 

With the final retreat of glaciation, du Toit 
(1921, p. 215) envisaged the development of " ... 
an immense estuary ... covering practically the 
whole area south-west of a line drawn from 
Mariental, past ... Vryburg to Port St. Johns; it 
connected with the ocean on the west and was 
presumably bounded by land on the south­
certainly on the north and east." Stratten (1970, 
p. 488) postulated two restricted openings to the 
ocean, one in the north-west near Loeriesfontein, 
the other in the south-east, near East London. 

The Douglas fauna provides the first clear 
evidence of marine conditions in the Great Karroo 
basin at that time. The marine character of the 
deposits at Blaauw Krantz gains further support 
from the occurrence of moulds of the mineral 
glauberite in concretions from the fossil-bearing 
horizons. David et al. (1905, p. 178) concluded 
that in Australia "The horizons ... being not far 
below ... the top of the highest beds of a Marine 
Series, where they are about to give place to fresh 
water beds ... , it is likely that the conditions 
favoured isolation of shallow sea basins ... ". 
Raggatt (1937, p.346) emphasised that cold 
marine conditions favoured the development of 
glauberite. He suggested that the decomposition of 
marine organisms might have provided the 
necessary sulphur. If the water were saturated with 
calcium carbonate, a slow warming, following the 
crystallisation of the glauberite, would induce 
precipitation of the carbonate. It would tend to 
accrete around a nucleus, perhaps a glauberite 
crystal or a cadaver, and so form concretions. 
Weeks (1953, p. 166) felt that for the formation of 
concretions, bottom conditions should be stagnant 
and anaerobic. The scarcity of bottom dwellers 
(brachiopods and lamellibranchs) at Blaauw Krantz 
supports this idea. 

According to Stratten (1970, p.488) "The 
Main area covered by this sea during the Dwyka 
Period is shown by the occurrence of the Upper 
Dwyka Shales" (see fig. 1). It is likely that the sea 
responsible for this fine-grained facies, which 
contains marine invertebrates at its base, extended 
farther north into the basin. This leads to 
speculation on the equivalent near-shore shallow­
water facies. The most obvious other indicator of 
marine conditions in the Great Karroo basin is the 
glauconite associated with the Coal Measures in the 

deltaic Middle Ecca sediments of the northern part 
of the basin. A marine origin for this glauconite has 
always been in doubt, as the fossil evidence has 
been sparse and ambiguous (McLachlan, 1973). It 
is not impossible that the shales of the Douglas 
fossil sites were accumulating in the deeper water 
of a shallow shelf sea (linked to the ocean in the 
west), while the glauconite was formed during brief 
local transgressions of the sea on to the brackish/ 
fresh-water coastal swamps where coal was 
accumulating i.e. that the White Band might be 
stratigraphically higher than the Middle Ecca coals. 
The few recorded "marine" fossils of the Ecca Coal 
Measures (sponge spicules, a problematic 
cephalopod, acritarchs) have seemed anomalous in 
a basin that lacked any typical marine macrofaunal 
assemblages (McLachlan, 1973). They too might 
have occurred in a marginal marine environment 
connected with brief transgressions of this sea. The 
facies distribution in the Kalahari basin is essen­
tially the same: marine Dwyka in the west and a 
deltaic sequence with Coal Measures in the east, 
which are traditionally considered to be younger 
(i.e. Ecca). Du Toit (1954, p.354) pointed out 
that in South America "The Rio Bonito group 
contains coal and has yielded an abundant flora ... 
strikingly like that of the Ecca series." Palyno­
logical studies may support this contention U. M. 
Anderson, in prep.). In the Great Karroo basin, the 
coals occur in the northern portion of the basin, 
and the White Band (demarcating the top of the 
Upper Dwyka shales) in the southern portion, and 
no single section has been found to include both. 
In South America, however, the coal lies below the 
Mesosaurus-bearing zone (White Band equivalent) 
(cf. du Toit, 1954, p. 355). This is also the case in 
South West Africa (see p. 37). 

The common occurrence of fossil wood at the 
Douglas sites certainly indicates a well established 
flora, including large trees, on the nearby land_ The 
well-rounded ends (see fig. 11) of many of the 
pieces of wood probably formed during stream 
transport down the Kaap valley to the site of their 
deposition, which may have been estuarine. 

White Band at the Top of the Upper Dwyka Shales 
Stage 

Venter (1969, p.15) and Stratten (1970, 
p.488) believed that the Upper Dwyka Shales 
accumulated under marine or estuarine conditions. 
Fossil evidence is lacking for the greater part of the 
shales, and that from the White Band is incon­
clusive: the fish Palaeoniscus capensis confirms 
only that the environment was aqueous, and the 
aquatic reptile Mesosaurus gives no indication of 
the salinity of the water. The crustaceans 
might, however, be cited in support of marine 
deposition as "There is no positive evidence that 
any of the eocarids actually inhabited fresh water" 
(Brooks, 1964, R337). They were benthonic 
creatures and " ... the crab-like No to caris , from 
the Permian of Africa, is the culmination of this 
adaptive trend." (op. cit.). The existence of this 



bottom fauna does suggest that the strongly 
reducing conditions inferred from the White Band 
lithology were confined to the sediment itself, and 
the overlying water was perhaps not markedly 
de-oxygenated. The absence of other benthonic 
organisms is puzzling. 

It is interesting that both the crustaceans and 
Mesosaurus have only been collected from the 
western side of the Great Karroo basin (fig. 1). The 
isopachs of the Upper Dwyka Shales show separate 
eastern and western sub-basins (Winter & Venter, 
1970, fig. 6), and it may be that conditions in the 
east were different at the time of deposition of the 
White Band. It is also likely that fossils have not 
been noted in the White Band east of Laingsburg 
because they have been obscured by the 
metamorphic effects of the Cape folding. A further 
point on the distribution of these fossils is their 
occurrence at positions ranging from near the zero 
isopach at Kimberley to the 600 ft. (183 m) 
isopach in the western Cape. 

Fossil wood and leaves have been recorded 
from several localities in the White Band 
(Haughton et al., 1953, p.23; du Toit, 1954, 
p. 279; Halbich, 1958, p. 124; Heath, 1966, p. 42). 
In the western Kalahari basin the White Band is 
separated from the marine faunas by nearly 350 m 
of sediment and an erosional unconformity (Heath, 
1966, PI. 1, p. 73), while the glauberite in the 
marine horizon at Blaauw Krantz indicates that 
conditions were becoming less saline from the base 
of the shales upwards (see above). The turbidite 
beds in the overlying Ecca sediments (Kuenen, 
1963, pp. 191-192; Ryan, 1967, p.953; 1968, 
p. 133; Theron, 1967; Truswell & . Ryan, 1969) 
have been quoted in support of a marine environ­
ment for the White Band (Meyerhof & Meyerhof, 
1972 p. 290), but the White Band itself is not 
known to contain turbidites and, furthermore, 
turbidity currents are not exclusively operative in 
saline water (cf. McBride, 1964, p. 94). 

CONCLUSION 

The fauna at Douglas confirms the existence 
of marine conditions in at least the western part of 
the Great Karroo basin and links it to the sea that 
occupied the Warmbad basin and the western 
Kalahari basin at the close of Dwyka glaciation. 
The marine invasion was apparently brief, and by 
White Band times conditions were possibly already 
non-marine, perhaps due to elimination of the 
connection with the ocean in the west. 

ACKNOWLEDGEMENTS 

We thank Dr. J. M. Dickins, Dr. C. Teichert, 
Dr. R. A. Jubb, Dr. M. E. Williams and Mr. J. C. 
Loock for specimen identifications, Mr. J. M. 
Anderson for allowing reference to his palyno­
logical data, and Dr. H. Martin for commenting on 
the geological section in the Kalahari basin in 
South West Africa. 

47 

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Symp. Pap, 1969, Interim report 35 pp. 
S.O.E.K.O.R. J ohannesburg. 

VISSER, D. J. L. (1958). The geology and mineral 
deposits of the Griquatown area, Cape 
Province. Geol. SUrD. S. A fr., Expl. Sheet 
185, 72 pp. 

VON HUENE, F. (1925). Die Sildafrikanische 
Karr oo-Formation als geologisches und 
faunistisches Lebensbild. F ortsch. Geol. 
Palaeont., 12; 124 pp. 

---- (1941). Osteologie und systematische 
Stellung des Mesosaurier. Palaeontographica 
92 A, 45-55. 

WASS, R. E. (1972). Permian bryozoa from South 
Africa. J. Paleont., 46; 871 - 873. 

WEEKS, L. G. (1953). Environment and mode of 
origin and facies relationships of carbonate 
concretions in shales. J. sedim. Petrol., 23 (3) , 
162-173. 

WILLIAMS, M. E. (1972). The origin of "Spiral 
Coprolites". Univ. Kansas Paleo. Contn·b. , 
Paper 59, 19 pp. 

WINTER, H. de la R. and VENTER, J. J. (1970). 
Lithostratigraphic correlation of recent deep 
boreholes in the Karroo-Cape sequence. 
LU.G.S. 2nd Gondwana Symp. , Froc. Pap. pp. 
395-408 South Africa 1970. Distributed by 
Geol. Soc. S. Afr. 

WOODS, H. (1922). Note on Pygocephalus from 
the Upper Dwyka shales of Kimberley. Trans. 
Geoi. Soc. S. Afr., 25; 41- 42. 



;a 
u ·60 .... 
o ·2 
0::> 

SITE 
\) 

c.!) 

Gaus and 
Itzawisis 

Gellap and 
s. Wasser 

Huns 

\) Viperstorf and s:: 
0 Tses .... 
'" "d 
;=j 

~ 
I Tses ... 

\) 

"d 
'3 
0 

p:) 

'" \) 

Brukkaros '" f-; 

Locali ty 
unspecified 

Between Asab 
and Tses 

Tses 

TABLE 1 

DWY KA INVE RTEBR ATE FAUNA- KALAHARI BASIN IN SOUTH WEST AFRICA 

For p lace names, see 1: 1,000,000 geological map of South West Africa (1963) . 

FOSSILS AV AILABLE SPECIMEN DETAILS SOURCE 

Lamellibranch: Numerous. 
Eurydesma cf globosum In dark shales above glacial tillite. Schroeder (1908, p. 695) identified specimens found by Range 
re-allocated to at I tzawisis. 
E. mytiloides by Range (1912, p. 29) recorded E. cf globosum from the farm 
Dickins (1961, p. 143) Itzawisis, and from Gaus, on the Kameelhaar river. 

Gastrop od, indet. Von Huene (1925, p. 27) listed Eurydesma and Conularia from 
Itzawisis and Gnus (? misquoting Range, 1912). 

Lam ellibranch: From turbidite sandstone interbedded with shale. Martin & Wilczewski (1970, p . 226) , name only. 
Eurydesma my tiloides 

Lamellibranch : 'steinkern ', isolated specimens Range (1912, p. 30). Tentative identification only. No figure. 
Inoceramus·like The position of this site, on Huns farm, 13 miles S.E . of 
bivalve Keetmanshoop, appears to fall within the Tses Boulder-

Gastropod , indet. mudstone (from Martin, 1953 , PI. 5) . 
Range (1912) records the fossils also from Gibeon. 

Gastrop od : Several hundred, but only 4 good enough to Dickins (1961) described and figured 4 specimens from the 
Peruvispira vipersdor- describe. farm Viperstorf 63 . He also mentioned 'other specimens' 
fensis In calcareous concretions. from 8 miles West of Tses siding, near the main road. 

(Geological Survey, Pretoria) 

Nautiloid cephalop od : Single fragmentary specimen in a concretion . Du Toit e915, p. xlv}, name only. 
Orthoceras sp. External and internal moulds of portion of Du Toit 1921, p . 214}, name only. 

phragmacone 5 em long. Haughton & Frommurze (1927, p. 142), brief description , no 
(Sou th African Museum, Cape Town, No. 8931.) illustration. 

Echinoid remains Fragmentary ambulacral plates and spines. Haughton & Frommurze (1927, p. 141), brief description, no 
Archaeocidaris sp . In a calcareous concretion. illustration, from W. of Brukkaros siding, near the main road. 

Echinoid remains: sp ines 
Sponge spicules In calcareous concretions. Martin & Wilczewski (1970, p. 226), listed only . 
Scolecondonts 

Crinoid columns In calcareous concretions. Martin (195 3, p . . 17), mentioned only. 
Gastrop od, indet. Martin & Wilczewski (1970, p. 226), mentioned only the 

crinoid columns. 

Foraminifera: In calcareous concretions, from 2 localities near Martin & Wilczewski (1970, pp. 226- 227) reported. Identifica-
genera Tses, close to one another. tions not yet completed. 
Hyperamm ina, 
A m modiscus, 
Glomosp ira, 
A m mobacculites, 
Spiroplectammina. 

'" o 



Schlip Cephalopod: Goniatite 
genus Eosianites, 
subgenus Glaphyrites. 

Radiolaria, indet. 

Hardap 110 Lamellibrach: 
Eurydesma my tiloides 

Gastropod, indet. 

Brachiopod, indet. 

~ 
Bryozoa, indet. Later 

~ identified as 
..c: Dyscritella cf. D. ct:J 

'" spinigera (Bassler) 
:.a 
0 

Asteroidean, family ..lI: 
'2 Monasteridae. os 
C.!l Genus and species 

indet. 

Asteroidean, indet. 

Mariental, E. of Lamellibranch: 
railway station Eurydesma globosum 

Ganikobis Gastropod, indet. 

Conularia sp. 

Radiolaria, indet. 

Locality unknown Brachiopod: 
Productus sp. 

- -- - -

A sin~le specimen, complete and well preserved. 
In a sIliceous, phosphatic concretion. 
(Geological Survey, Windhoek) 

In siliceous, phosphatic concretions. 

At least 5 moderately well-preserved specimens. 
In sandstone turbidite beds. 
(Geological Survey, Pretoria) 

One specimen. 
In fine grained sandstone. 
With bryozoan fragments. 

In sandstone turbidites, intercalated in shales. 
Identified by Dr. Glaessner. 

In sandstone turbidites, intercalated in shales. 

Single arm only. 
Associated with Eurydesma mytiloides 
and bryozoan fragments. 
(United States National Museum collections. 

No. 16259). 

Occur in sandstone turbidites interbedded with 
shale. 

According to Martin (pers. comm.) the specimens 
(in the Windhoek Museum) have been lost. 

In sandstone and shale, together with worm 
burrows and Phyllotheca-like stems. 

Single specimen. 
In siliceous concretion. 

Single specimen. 
In siliceous concretion. 

In siliceous concretions. 

No details. 

Martin, Walliser & Wilczewski (1970), described and figured 
(line drawing). 

Martin, Walliser & Wilczewski (1970, p. 621) , mentioned only . 

Martin (1953, p. 37), mentioned only. 
Dickins (1961) described and illustrated. 

Wass (1972, p. 871), mentioned only. 

Heath (1966, p . 30, PI. XVII). 

Martin & Wilczewski (1970, p. 226), listed. 
Dickins (1961, p. 138). Mentioned only. 
Wass (1972), described, illustrated, named. 

Lane & Frakes (1970) . In the Fish River valley, a few km 
N.W. of Mariental i.e. within 10 km of Hardap 110. 

Martin & Wilczewski (1970), listed one. 
Heath (1966, p . 30, PI. XVIII) found a specimen in his " yellow-

green mudstone" zone. 

Du Toit (1916, p. 11), mentioned only. 
Heath (1966, p. 30) concluded that they derived from the same 

zone as the fish at Ganikobis i.e. Ganikobis shale. 

Martin & Wilczewski (1970 , p. 226) , listed. 

Schroeder (1908, p. 697) reported. 
Range (1912, Fig. 3), illustrated. 

Martin & Wilczewski (1970, p . 226), listed. 

Von Huene (1925, p. 27) noted a Productus in Dr. Reuning's 
collection. 

'" ...... 



-; 
u 

·So .... 
o·~ SITE - I:: o::;J 
\I 
(j 

Haib 

II) 

-; 
-= Kanibeam en 

'" ..10: 
:>-
~ 
0 .. 

Klipneus II) 

0. 
0. 

::;J .... 
0 
.... .. 
'" 0. .. 
II) 

~ 
0 

....l 
Zwartbas 

TABLE 2 

DWYKA INVERTEBRATE FAUNA-WARMBAD BASIN, SOUTHERN SOUTH WEST AFRICA 

For location of sites, see 1: 250,000 geological sheet Amib (H-33-F) in Haughton & Frommurze (1936). 

FOSSILS AVAILABLE SPECIMEN DETAILS SOURCE 

Lamellibranch: 2 specimens, one being very poor and fragmentary. Reed (1935), described and figured. 
Aphanaia haibensis (Probably housed in the South African Museum, 

Cape Town. (S.A.M.}.) 

Gastropod, indet. 1 specimen, a coiled shell, too poor for any Reed (1935, p. 161), discussed. No. illustration. 
satisfactory identification. 

(Probably S.A.M.) 

Arenaceous foraminifera: In calcareous concretions. Extracted by acid Martin & Wilczewski (1970, p. 227) . Mentioned only. 
almost identical with treatment. No further information. 
those from Tses, (Martin 1971, written comm.) 
Kalahari basin. 

Lamellibranch, indet. 2 poor in ternal moulds, in a calcareous 
concretion. 

(Bernard Price Institute for Palaeontological 
Research (B.P.I.P.R.) 1.41.) 

Arenaceous foraminifera In calcareous concretions from fossil fish horilon. 
Radiolaria Extracted by acid treatment. Recently collected by the authors . 
Sponge spicules. (B.P.I.P.R.) 

Investigation proceeding. 
Lamellibranch, indet. 1 specimen, external mould, in a calcareous 

concretion. 
(B.P.I.P.R. 1.43) 

Arenaceous foraminifera. 4 specimens visible in 2 calcareous concretions 
also containing fish. 

(B.P.I.P.R. P.2 and P.6) 

Heavy lines indicate new finds. 
- -

(.Jl 

r...o 



-; 
u 

·60 .... 
0·2 
o~ ., 
o 

'0 
§ 
~ ., 

~ 

... '" ., ., ~ ., 
o.?-~ 
o.~..c: 
~O{/) 

TABLE 3 

DWYKA INVERTEBRATE FAUNA-GREAT KARROO BASIN, SOUTH AFRICA 

SITE 

Kimberley area 

FOSSILS 

Crustacean: 
subclass: Eumalacostraca 
superorder: Eocarida 
order: Pygocephalomorpha 
family: Notocarididae 

Notocaris tapscotti 

Loeriesfontein Crustacean: 
area family: Notocarididae 

Laingsburg Crustacean: 
family: Notocarididae 

Py gasp is ginsburgi 

AVAILABLE SPECIMEN DETAILS 

Numerous specimens. 
Impressions in weathered White Band. 
(A.M.M.M. * and Albany Museum, Grahamstown.) 

Numerous specimens. 
Impressions in weathered White Band. 
(S.A.M.*, B.P.I.P.R.* and Stellenbosch 

University. ) 

Numerous specimens. 
Impressions in White Band. 
(S.A.M. and B.P.I.P.R.) 

SOURCE 

Rogers & du Toit (1909, p. 193): "probably Anthrapalaemon 
sp.", listed from the Upper shales. 

Haughton (1919, p. 9) " ... provisionally assigned to the 
Carboniferous genus Anthrapalaemon by Reed, in the 
White Band at the boring at Elandsdraai, Orange River 
Station". 

Woods (1922) described and figuredPygocephalus sp., from 
near Kimberley. 

Broom (1931) described and figured better material from 
Woods' (1922) site, and renamed the fossil No to caris 
tapsc0 tti. 

du Toit (1954, p. 279) listed "Notocaris (Pygocephalus) 
tapscotti Broom, Kimberley, and A nthrapalaem on sp., 
Orange River Station" (near Hopetown) from the White 
Band. 

Haughton (1969, p. 355): "From Kimberley ... Notocaris 
tapscotti and from Orange River Station another Crustacean 
Anthrapalaemon sp." Pygaspis ginsburgi Fabre not 
mentioned. 

Brooks (1969, p. R341): "Anthrapalaemon is believed to be a 
synonym of Pygocephalus" . 

We conclude, therefore, that the names Anthrapalaemon sp., 
Pygocephalus sp. and No to caris tapscotti probably refer to 
identical crustaceans from the White Band in the 
Kimberley- Orange River Station vicinity, and that 
Notocaris tapscotti is the only valid name. 

Its relationship to Pygaspis ginsburgi from the White Band at 
Laingsburg has not yet been established by direct 
comparison. 

Recently collected from the farm Ezels Fontein. 

Fabre (1967), described and figured. He did not re·examine 
Notocaris tapscotti. 

Small crustaceans also occur in the Iraty Formation in Brazil. The Iraty Formation is considered to be the equivalent of the White Band (du Toit, 1954, p. 353). 
Bigarella and Salamuni (1967, pp. 64- 65) list Pygaspis braziliensis, P. quadrata, Paulocaris pachecoi, Clarkecaris brazilicus, Liocaris huenei and L. augusta. 

Sambokkraal 
borehole 

Sponge? spicules Haughton et al. (1953, pp. 22, 40) mentioned "spicules, 
presumably from sponges" from a calcareous, carbonaceous 
shale, some 110 feet (34 m) below the base of the White 
Band. 

~ 

*" 



.s ~ Matjiesfontein Radiolaria 5 specimens figured from a single thin section of Strydom (1964, p. 279, PI. II), briefly described and 

.... [~ phosphorite rock (Slide stored at Stellenbosch illustrated. 
o p...: University, Geology Department. Apparently Judging from the photographs and descriptions, it is possible 
~;:J ~ now lost.) that the "radiolaria" migh t, in fact, have been spinose 

l:Q 0 acritarchs. 

Laingsburg Radiolarian spicules "Probable radiolarian spicules occur in the cherty A. C. Theron (1967, pp. 15,22), mentioned. (See forgoing 
II) textured part of the graded interval" of the quote.) 

... ~ Matjiesfontein chert member, which is a Both Mr. A. C. Theron and his M.Sc. supervisor Prof. I. C. 
I:l ~ turbidite sequence. It lies within the Lower Rust (1972, written comms.) stress that the identification 
-tl c<I Ecca series, a few feet above the White Band. was extremely tentative . 
• 1:: e A. C. Theron (1967, p.15) erroneously referred Strydom's 
B ~ (1964) "radiolaria" to his Lower Ecca Zoutkloof shale 
§ I:l member, immediately above the White Band. 
't: ~ Teichert (1970, p.132):" ... the 'White Band', which 
:~o-l contains Radiolaria and is overlain by a radiolarian chert 
~ ~ U. N. Theron, pers. comm.)". Dr.J. N. Theron (1972, 
:21.. written comm.) states that Teichert (1970) misquoted him . 

. S Rust and A. C. Theron (1972, written comm.) confirm that 
there has been no report of Radiolaria in the White Band. 

Lamellibranchs: 
(i) Nuculid? 1 complete specimen, internal and external 
? Nuculopsis sp. moulds, in a calcareous concretion. Recently collected by the authors. 

(B.P.I.P.R. 1.39) (see figs 22, 23 ). Dr. J. M. Dickins made identifications from stereopair 

(ii) Nuculanid Complete specimen, internal and external moulds, 
photographs. 

Phestia sp. and 2 other partial external moulds, all in a single 
II) calcareous concretion. 
~ (B.P.I.P.R. 1.38) (see figs 24, 25, 26) . 
~ II) ----~------------4----------------------------------------------------
~ -Q, Brachiopod: Several specimens, internal moulds, all in a single Recently collected by the authors . 
....: ;::I Attenuatella sp. calcareous concretion. Dr.J. M. Dickins made the identitification from the specimens 
~ 8 (B.P.I.P.R.1.40). themselves. 
o ~ ----~-------------------------------------------

I:l 11.1 Orthocerid cephalopods: 
p.. :; (i) 8 specimens in calcareous concretions. Most are 
:§' ~ external moulds, 2 of which are nearly 

11.1 .;;; complete. 2 specimens show internal and 
-5 ~ external moulds of the body chamber (figs. 
'0 r-- 12,13,14). Septa are occasionally present. 
~ The siphuncle is central. 
~ (B.P.I.P.R. 1.1 - 1.8). 

One incomplete specimen was found at Zand 
Bult. External and internal mould of part of Recently collected by the au thors. 
phragmacone. Identification confirmed by Dr. C. Teichert, from stereopair 

photographs. 
(ii) 1 incomplete specimen in a calcareous concretion. 

External mould of portion of phragmacone. 
Septa and siphuncle partially preserved. 

(B.P.I.P.R. 1.9) (see fig. 16). 

* A.M.M.M. = Alexander McGregor Memorial Museum, Kimberley. 
S.A.M. = South African Museum, Cape Town. 
B.P.I.P.R. = Bernard Price Institute for Palaeontological Research, Johannesburg. 

Heavy lines indicate new finds. 

(J1 
(J1 



~ 
Z u - ·SO .... 
V) o·~ SITE -( -I: 
I=Q ~;:J 

C!> 

Schlip 

Z Ganikobis - IlJ V) 
~ -( 

I=Q ..c: 
V) - '" ~ :E -( 0 

::t:: ....: 
-( '2 
...:I '" 
~ 

C!> 

Haib 

'" IlJ 

~ 
Z ..c: - V) 

Viool's Drift V) «I 
-( ....: 
I=Q >-

~ 
Cl Cl 
-( ... Klipneus** I=Q IlJ 

::E p. 
p. 

~ ;:J 
-( .... :: 0 

IlJ 

'" Zwartbas** «I 
I=Q 

Ezels Fontein, 
farm near 
Loeriesfontein 

TABLE 4 

DWYKA FISH FROM THE KALAHARI, WARMBAD AND GREAT KARROO BASINS 

FOSSILS AVAILABLE SPECIMEN DETAILS SOURCE 

Palaeoniscoid fish, indet. In siliceous concretions Martin & Wilczewski (1970) . Mention only. 

Palaeoniscoid fish: 
(a) Namaichthys schroederi 7 specimens in calcareous concretions. Giirich (1923) described and illustrated Namaichthy s 

Holotype lost. schroederi. 
(N eotype at Geological Survey, Pretoria, Gardiner (1962) re-described and figured new material. 

others at British Museum of Natural 
History.) 

(b) Watso'nichthys lotzi 1 specimen of incomplete fish. Giirich (1923) described and illustrated Acrolepis lotz i. 
(Original material at Central Geological Gardiner (1962) renamed. 

Service, Berlin. Presumed lost.) 

(c) Elonich th y s? 

~d) Rhadinich thy s? Giirich (1923) described and illustrated. 
e) Genus V 

Fish scales, indet. In concretions. Haughton & Frommurze (1927, p . 141). Mention only. 
Also a "fragment of a large amphibian (? ) 

bone". No further details. 

Palaeoniscoid fish fragments In concretions. Haughton & Frommurze (1927, p. 141) . Mention only. 
"assigned to the genus 
Acrolepis". 

Palaeoniscoid fish, indet. 3 specimens in calcareous concretions and a Recently collected by the authors. 
number of isolated scales in individual 
concretions. 

(B.P.I.P.R.* P.7-P.I0.) 

Palaeoniscoid fish: 
Namaichthys schroederi? 6 specimens in calcareous concretions. Dr. R. A. Jubb made identifaction from specimens recently 

(B.P.I.P.R. P.I-P.6.) collected by the authors. 

Palaeoniscoid fish, indet. Several specimens. Impressions only. B.P.I.P.R. specimens collected recently. 
(S.A.M. *, Stellenbosch University, B.P.I.P.R.) Dr. R. A. Jubb likened the fossils to one of the incomplete 

specimens (S.A.M. 1066) described by Broom (1913a) as 
P. cap ensis. See below. 

-----_. -

(j1 

c:r> 



Hantamsberg Palaeoniscoid fish: 3 incomplete specimens, none having a head. Rogers & du Toit (1909, p . 193 ) " White Band . . . in 
on farm Toren Palaeoniscus capensis Impressions in shale. Calvinia .. . Elonich thys sp." 

Z near Calvinia Broom (1913) said they were "probably Broom (1913 a) described and figured P. capensis from 12 - Upper Dwyka". He did not mention the miles west of Calvinia. No mention of Elonich th ys sp. rn 
<: discoverer. from Calvinia. 
~ Roger & du Toit (1909) listed Elonichthys du Toit (1954 , p.279) "White Band- Palaeoniscus capensis 
0 "t:l sp . from the White Band near Calvinia. Broom, Calvinia". No further mention of Elonich th ys. 
0 I: du Toit (1954) listedP. capensis, Broom Gardiner (1962, p.18) : "Elonich thys sp., from the ~ <'d 

~ 
a:I from the White Band, with no mention of White Beds of the Upper Dwyka shales, * Clavinia" 
II) <: 

~ 
Elonichthys. (*Typing error? Should read Calvinia" ). 

~ We gather that du Toit (or Rogers) was the We conclude that Broom (1913a) possibly renamed 
E-< 
<: finder of the fossils near Calvinia, and Elonichthys sp. from Calvinia as Palaeoniscus capensis. 
~ that they derive from the White Band. If correct, the latter is the valid name. 
~ (S.A.M. 1061,1062,1066). Dr. R. A.jubb (1972, written comm.), suggests that the 0 three specimens of P. cap ensis described by Broom, 

probably represent more than one species. 

Langb erg Palaeoniscoid fish " .. . undoubted fin- and scale-impressions of Hiilbich (1958, p.124) . 
mountain, Paleonisclls were seen." 
ab out 20 
miles N.W. of 
Loeriesfontein 

". Kimberley, Palaeoniscoid fish: 5 fragmentary specimens on a single shaly Broom (1913b) described and illustrated. <'d 
u Diamond Acrolepis addamsi sandstone block displaced within a Dr. R. A.jubb (written comm., 1972) : may involve 2 genera u 
~ pipe Kimberlite pipe. Horizon, therefore, 
". cannot be ascertained with confidence. <'d 
~ Broom said it might be either Dwyka or >-
~ Ecca. (We feel it could also be Beaufort.) 

Cl (A.M.M.M. ) 

'0 ... ~ III 
Blaauw Palaeoniscoid fish: Sev~ral specimens of dismembered fish in Identified by Dr. R. A. jubb from A.M.M.M. specimens. 

II) II) >-~ Kran tz** Namaichthys schroederi? calcareous concretions. B.P.I.P.R. specimens recently collected by the authors. 
III Q.. ~ <'d and oth ers (A.M.M.M., collections 4902,5008; ~&Cl~ B.P.I.P .R. P.ll - P.23 , P.29- P.42.) 

----

* B.P.I.P.R. = Bernard Price Institute for Palaeontological Research , Johannesburg. 
S.A.M. = South African Museum, Cape Town. 
A.M.M.M. = Alexander McGregor Memorial Museum, Kimberley . 

**Coprolit es were found by the au thors at Klipneus (B.P.I.P.R. 1.42), Zwartbas (B.P.I.P.R. 1.44) and Blaauw Krantz (B.P.1.P.R. 1.18- 1.3 7) (Figs 17- 2 1). There are true 
coprolites as well as occasional h eteropolar "spiral cop rolites" that probably represent fossil spiral valves from the intestines of chondrichthyians (Williams, 1973, 
written comm.) . 

Heavy lines indicate new finds. 

(J1 ...., 



TABLE 5 

LITERATURE ON THE MESOSAURIDAE FROM SOUTHERN AFRICA. 

AUTHOR IDENTIFICATION SITE MUSEUM NO. FOSSIL DESCRIPTION 

Gervais (1865) Mesosaurus tenuidens Griqualand West , north of Museum of Natural Head, neck, forelimbs and most 
the Orange River History, Paris of thoracic and abdominal 
(Kimberley area). region. 

Giirich (1889) Ditrich osaurus Near Hopetown ? Posterior region, with 1 slightly 
capensis imperfect forelimb, ribs and 

vertebrae 

Seeley (1892) Mesosaurus Y<I ml. S. of Market British Museum Ribs and vertebrae. 
pleurogaster Square, Kimberley. (B.M.) 49972 

-
" 

100' from E. margin B.M.49971 Ribs and vertebrae. 
of Kimberley mine. 

" 
40' from E. margin B.M.49973 Vertebrae and ribs. Seeley 
of Kimberley mine. suggested it migh t be part of 

B.M.49974. 

" " 
B.M. 49974. Vertebrae, ribs and hind foot. 

Mesosaurus tenuidens Albania, Griqualand S.A.M. ? 709 Head, forelimbs, torso. 
West. Ventral aspect. 

Mesosaurus sp. Near Burghersdorp. Albany Museum, Dorsal aspect of vertebrae and 
Grahamstown. ribs. 

Broom (1904) Probably Discovered in Bushman- No mention. Most of tail, hind limbs, 
Mesosaurus tenuidens land by the Rev. Neethling abdominal and thoracic region 

of NieuwoudtviIle and portion of skull. 

Broom (1908, 1. Mesosaurus Near Calvinia. S.A.M. ? 9327 Cervical region, ribs, left 
p. 379) tenuidens forelimb, poor skull cast. 

Gervais 

---

COMMENTS 

Described only. No iIlustration. Found in 
about 1844 in Griqualand, north of the 
Orange River (i.e. in the Kimberley area). 
The specimen slab was being used by a 
Hottentot as a pot·lid! 

Described and figured. 
"Remains are evidently those of an immature 

animal, and there seems little doubt that 
they are those of a young M esosaurlls" 
(Broom 1904 p.103). 

Described. 

Described. 

Described and figured. 

Described. 

Described and figured. 
Albania is a former farming area in the 
magisterial district of Herbert, between 
Hopetown and Douglas. 

Brief description, no iIlustration. 
The town lies on Beaufort sediments close 
to the Stormberg contact, nearly 200 miles 
(320 km) from the nearest Dwyka outcrop. 
This record is suspect, as it is stratigraphicall; 
incompatible with other Mesosaurus 
records. 

Described and iIlustrated. 

Described and iIlustrated. 
"Species represented by the Paris type 
fGervais 1895), this Calvinia specimen 
Broom 1908), and possibly also the 

Nieuwoudville specimen {Broom 1904)". 

(.;>l 
ex> 



2. Mesosaurus 
capensis 
(Giirich) 

3. Mesosaurus 
pleurogaster 
Seeley 

Broom (1913c, Noteosaurus 
p.358-360) africanus 

Reuning & Mesosaurus tenuidens 
von Huene 
(1925 ) 

von Huene Mesosaurus tenuidens 
(1925) 

von Huene 
(1941, p.55) 

On the S.A.M. specimen described by Seeley 
(1892): "The specimen .... must thus be 

• regarded as a distinct species" (from 
M. tenuidens). 
Comparing Giirich's (1889) type specimen 
and the S.A.M. specimen described by 
Seeley (1892), Broom concluded that it is 
"highly probable that the Cape specimen is 
the adult of Giirich's type". 
"M. capensis represented by Giirich 's 
type and the specimen described by 
Seeley from the S.A.M." 

"Very imperfectly known". 

Victoria West S.A.M. 2355 Pelvic region with the anterior Described and illustrated. 
district part of the tail and the "the possession of 6 phalanges in the 5th 

greater part of the two hind toe of Noteosaurus is a striking difference 
limbs. from Mesosaurus when certainly there are 

only 4 in the known specimens". 
With regard to its geological setting: " There 
is no doubt it is from the same horizon as 
the South African species of Mesosaurus". 

Haughton (1920, p . 9) , however, noted that 
in no portion of the Victoria West district 
has the White Band been mapped, although 
it occurs just outside the boundary to the 
N. of Vosburg. He concluded that "It is 
just possible, therefore , that Noteosaurus 
is from the Lower Ecca shales" . 

We regard the location of Broom's 
Noteosaurus as effectively unknown. 

Doros, in the ? Fractured, disarticulated ribs, Originally described as occurring in a 
Kaokoveld. vertebrae and some extremity volcanic breccia underlain by tuffaceous 
20° 45' S. 14° 15' E. bones. No.skulls. beds containing volcanic "Kalkbomben". 

Martin (1961, pAO) re-examined the site 
and described the fossil bed as being a dark 
course , conglomeratic quartzite band about 
1 foot thick containing numerous rolled 
and broken Mesosaurus bones. The 
erroneously described "Kalkbomben" of 
Reuning are apparently calcareous 
concretions. Coaly carbonaceous shales 
occur about 300 ft (90 m) below the 
Mesosaurus zone and seem to be equivalent 
to the Auob sandstone coals. 

He reviewed the literature without re-examination of the South African material, and suggested that all the previously described 
South African Mesosaurus species might be included under M. tenuidens. He made no mention of N oteosaurus africanus. 
In discussing the dispersed bones in the Doros bonebed in the Kaokoveld (north of the Windhoek highlands), he concluded that, 
because of the lack of distinguishing features, they belong to the same species as occurs in the Kalahari and Karroo basins, 
namely M. tenuidens. 

With regard to Noteosaurus: "I suspect that the data given on Mesosaurus (including Stereostemum etc.) of 4 phalanges in the 
5th toe is based only on incomplete preservation. If this is so, the genus name N oteosaurus has to disappear". 

continued overleaf 

(.Jl 

<.0 



TABLE 5- Continued 

AUTHOR IDENTIFICATION SITE MUSEUM NO. FOSSIL DESCRIPTION COMMENTS 

Haughton & Mesosaurus sp. Kanibeam, Warm bad No mention "A few moulds of bones" Only mentioned. 
Frommurze basin. 
(1927, p. 14) 

Hau1hton & M. tenuidens Bibliographic list only. 
Brin (1954) (Gervais) von Huene (1925,1941) was not mentioned. 

M. pleurogaster 
(Seeley) 

M. capensis 
(Giirich) 
Broom 

Noteosaurus 
africanus 

- ,-

Stromer (1914) Mesosaurus sp. Kabus (Khabus 146), ? A hind limb and a hand. Described and illustrated. 
farm N. of 
Keetmanshoop, S.W.A. 

Martin (1953, Mesosaurus sp. Gellap-Ost 3 and Windhoek Museum? ? Localities shown on map. 
PI. V, p. 39) Khabus 146, farms N. of The Mesosaurus remains are from a "fine, 

Keetmanshoop, S.W.A. hard siltstone .... The White Band, which 
can still be recognised in the Warmbad 
district, is not developed here". 

von H uene (19 25) referred to the Khabus 
specimen as M. tenuidens. 

Martin (1961, Mesosaurus sp. Gross Daberas, 17 miles Windhoek Museum P.C. 37*: lower part of body Only mentioned. 
p.14). N.E. of Tses, S.W.A. and portion of tail. Poorly *Heath (1966, PI. XXII) illustrated specimen 

preserved. P.C.37. 
c. ------

There are undescribed specimens in various museum collections (their localities are indicated on fig. 1): 
1. In S.A.M. (South African Museum, Cape Town) there are about ten specimens (without skulls) from sites in the Western Cape: Laingsburg gypsum works; Tankwa Karroo, 

N.W. of Laingsburg; Toren, 12 miles (19 km) W. of Calvinia; Ezels Fontein farm, N.W. of Loeriesfontein; Brandvlei, 120 km E.N.E. of Loeriesfontein. 
2. Mr. B. Oelofsen of Stellenbosch University Zoology Department is presently making a collection from the Loeriesfontein district. 
3. The A.M.M.M. (Alexander McGregor Memorial Museum, Kimberley) has rib, vertebrae and limb moulds, mostly from the Kimberley townlands, but there are also specimens 

catalogued from farther afield at Belmont, Heuningneskloof and Kameelpan (32 km N.E. of Kimberley). 
4. The State Museum in Windhoek, S.W.A. apparently has a number of specimens. We have confirmation of one from near Haib (Mr. C. G. Coetzee, Director, written comm. 

to Dr. E. P. Plumstead, B.P.I.P .R.),? three (with skulls) from Kirchberg farm, N .W. of Warmbad, and? one from Commissioner's Pan, N .E. of Loeriesfontein (Dr. A. Keyser, 
Geological Survey, Pretoria, pers. comm.). 

It would seem that the Mesosauridae are in need of revision before meaningful comparisons can be made between the African and South American species, where M. braz iliensis 
and Stereosternum tumidum occur. 

0'> 
o 



Fig. 6: 
Fig. 7: 

Fig. 8: 
Fig. 9: 

Fig. 10: 
Fig. 11: 

Grey shale with bands of fossiliferous concretions 

Blaauw Krantz Site viewed from the east, showing stratigraphy. 
Varved shale at Blaauw Krantz. Dropstones to the right and left of hammer 
head, arrowed. 
Concretion in situ at Blaauw Krantz. 
Fossiliferous zone at Blaauw Krantz. "Q" indicates Quaternary sediments. The 
arrows indicate fossiliferous concretion horizons. 
Fossil wood from Blaauw Krantz (Specimen BK3). Transverse section (XY2). 
In situ fossil wood at Olie Rivier. Note rounded end. 

61 



62 

Orthocerid cephalopods-Blaauw Krantz 
Figs. -12-l4: Internal moulds of body chamber (xl). 

Fig. l2-Lateral view 1.3 (Stereopair xl). 
Fig. l3-Dorsal view 1.3 (xl). 
Fig. 14-Ventral view 1.3 (Stereopair xl). 

Fig. 15: External mould, showing ornamentation 1.4 (x2). 



Blaauw Krantz fauna 
Fig. 16: Orthocerid cephalopod. External cast and partly preserved septa and siphuncle, 

1.9 (Stereopair xl). 
Fig. 17: Coprolite, consisting partly of fish scales P.44 (XY2). 
Figs. 18-21: "Spiral coprolites" 

Fig. 18-Longitudinal section of heteropolar spiral coprolite ("enterospira" of 
Williams, 1972) 1.20 (Stereopair xl). 
Fig. 19-Longitudinal section showing more coarsely coiled structure 1.24 (xl). 
Fig. 20-Transverse section of coprolite with finely coiled structure 1.18 (x2). 
Fig. 21-Transverse section of more coarsely coiled coprolite 1.26 (x2). 

63 



64 

Blaauw Krantz fauna 
Figs. 22-26: Lamellibranchs (x5). 

Fig. 22-Nuculopsis sp. External mould. Lateral view, 1.39 (Stereopair). 
Fig. 23-Nuculopsis sp. Internal mould. Lateral view, 1.39 (Stereopair). 
Fig. 24-Phestia sp. External mould. Dorsal surface 1.38 (Stereopair). 
Fig. 25-Phestia sp. Internal mould. Dorsal view 1.38 (Stereopair). 
Fig. 26-Phestia sp. Internal mould. Lateral view 1.38. 

Fig. 27: Palaeoniscoid fish skull in concretion. Lateral view P.15 (xl).