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ANNALS OF THE ANNALE VAN DIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

VOLUME, 95 BAND 95

ie 7 ey) i :

*, ° 7 My

r one] t vps tt : f

" 2 r i

1 F *

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(

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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

VOLUME 95 BAND

ik VRUSTEES OF THE DIET RRUSREES, VAN GDIE

SOUTH AFRICAN MUSEUM SUID-AFRIKAANSE MUSEUM

CAPE TOWN KAAPSTAD

1985

EIS ORF CONTENTS

Page

Cooper, M. R.

A revision of the ornithischian dinosaur Kangnasaurus coetzeei Haughton, with a

elassinicanonoLine Ormithischiay (Rublishedtiuned9Ss.) ers eee ee 281

GosLiner, T. M.

The aeolid nudibranch family Aeolidiidae (Gastropoda, Opisthobranchia) from

MmoOpicalsouuneneAuricas (kublishedunel9 85>) many eee eer 233

GRIFFITHS, R. J.

Description of a new South African arminacean and the proposed re-instatement of

the genus Atthila Bergh (Mollusca, Opisthobranchia). (Published June 1985.)... 269

KENNEDY, W. J. & KLINGER, H. C.

Cretaceous faunas from Zululand and Natal, South Africa. The ammonite family

Kossmaticeratdae Spath> 1922° (Publishedtiune 1985.) ey = aes esas ee 165

KENSLEY, B.

The faunal deposits of a Late Pleistocene raised beach at Milnerton, Cape Province,

SoOumpaunicas(aublishedvApnill985s)) 2 te 455s ye eee ee eee oe ltt

KLINGER, H. C. see KENNEDY, W. J.

Orson, Si L.

Early Pliocene Procellariiformes (Aves) from Langebaanweg, south-western Cape

BIOMiNcee SOUUNEAtICaa(eublishedrApmil 1985.) as... one ase ee] ane is coe 123

Otson, S. L.

An early Pliocene marine avifauna from Duinefontein, Cape Province, South Africa.

(LPUULDNS OSG! ZA OTT USCIS) ee aoe ere ern ene eR OE Oo ann Ae) ee esc) Gea 147

SCHOLTZ, A.

The palynology of the upper lacustrine sediments of the Arnot Pipe, Banke,

Namaqualand (rublishedtAvornllOSs:). 55.4545. 5e meen ore ee ie 1

Volume 95 is complete in 8 parts.

VOLUME 95 PART 1 APRIL 1985 ISSN 0303-2515

. |e

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BULLOUGH, W. S. 1960. Practical invertebrate anatomy. 2nd ed. London: Macmillan.

FISCHER, P.—H. 1948. Données sur la résistance et de le vitalité des mollusques. J. Conch., Paris 88: 100-140.

FiscHER, P.-H., DuvAL, M. & Rarry, A. 1933. Etudes sur les échanges respiratoires des littorines. Archs

Zool. exp. gén. 74: 627-634.

Konn, A. J. 1960a. Ecological notes on Conus (Mollusca: Gastropoda) in the Trincomalee region of Ceylon.

Ann, Mag. nat. Hist. (13) 2: 309-320.

Konn, A. J. 19606. Spawning behaviour, egg masses and larval development in Conus from the Indian Ocean.

Bull. Bingham oceanogr. Coll. 17 (4): 1-51.

THIELE, J. 1910. Mollusca: B. Polyplacophora, Gastropoda marina, Bivalvia. In: SCHULTZE, L. Zoologische

und anthropologische Ergebnisse einer Forschungsreise im westlichen und zentralen Siid-Afrika 4: 269-270.

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(continued inside back cover)

ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 95 Band

April 1985 April

Part 1 Deel

THE PALYNOLOGY OF THE

UPPER LACUSTRINE SEDIMENTS OF THE

ARNOT PIPE, BANKE, NAMAQUALAND

A. SCHOLTZ

Cape Town Kaapstad

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THE PALYNOLOGY OF THE UPPER LACUSTRINE SEDIMENTS

OF THE ARNOT PIPE, BANKE, NAMAQUALAND

A. SCHOLTZ

Department of Archaeology, University of Stellenbosch

(With 21 figures and 2 tables)

[MS accepted 7 December 1983]

ABSTRACT

The Arnot Pipe on the farm Banke, Namaqualand, is one of the most southerly in the

Gamoep cluster of volcanics, which include ‘kimberlite’ and olivine-melilitite volcanic pipes. The

upper sediments of this pipe, in which palynomorphs as well as plant macrofossils and vertebrate

remains are found, are of lacustrine origin, having been laid down in a small, deep crater lake

formed in a vent of a ‘kimberlite’ volcano. Radiometric dates from other pipes in. the. cluster

indicate that the volcanic activity occurred between 64 and 71 Ma ago. In this study the systematic

palynology of the top 20 m of polleniferous sediments is described.

Seventy-two forms of spores, conifer and angiosperm pollen are described and illustrated

and 64 are formally classified. Fourteen new species and one new genus are defined and

21 further forms are, as far as is known, undescribed in previous literature. The possible affinity

of the fossil forms to 28 plant families, including, amongst the angiosperms, the Proteaceae,

Restionaceae, Ericaceae, Epacridaceae, Euphorbiaceae, Thymelaeaceae, Chloranthaceae,

Casuarinaceae, Cornaceae, Caesalpinaceae and Anacardiaceae, are suggested. The relevance of

these observations to hypotheses about the nature of Palaeogene vegetation in the African

subcontinent are discussed. The Banke palynoflora is contrasted with those of a similar time

range from tropical Africa and Australia to highlight what is unique about the combination of

taxa present in an early Tertiary vegetation in southern Africa. Particular attention is paid to

discussing the early history of some taxa that are at present characteristic elements of the

Capensis Flora.

CONTENTS

PAGE

MITER OCU HOM ae aes eae toi nd CN pe Ny hen gat ale Lahn 2

The location and geology of the Arnot Pipe ................ 3

Previous studies of fossil material from the Arnot Pipe....... 7

DY AGIIN ES Mapp eee eh, cee eect ae ore hig No meetreen U8 ace net eas 9

Waterialtandumethodsieio ae, ane nmeege cides so en ere 10

Rr ANOL OD Weg rere is eA Ce ate See ela, echt ca boe aie eae eat 13

Species list and register of type specimenms.............. 14

IDESCHIPTOMS Aon A Rie y. oN eR ee areal aes Lette cg ae 7)

OME Mi COUMES tvs syaeens nce eesti vet curt gtie eee le ees 88

DISCUSSION set epee eta ht he. eet flies oer ater « tee tare me tt tuegn al eee aw! 88

PNCKNOWICUSEMEMUSi te ie ot ee See e aa coed 103

NCTC TENCE Sr mibiar ney fe ential Rd hc: 5 MRS ky MAN ag ann leh einen 103

Ann, S. Afr. Mus. 95 (1), 1985: 1-109, 21 figs, 2 tables.

D ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

This paper presents the first detailed account of an early Tertiary

palynological assemblage from the subcontinent of southern Africa. To date,

direct fossil evidence relating to late Cretaceous and Tertiary vegetation history

and the evolution of plant groups in the region is very limited. The few relevant

palynological studies are mentioned below.

Cursory studies have been published on the Arnot Pipe (Kirchheimer 1934),

on the Knysna lignites—which may relate to a brief period in the Neogene

(Thiergart et al. 1963)—and on a DSDP sequence of mid- to late Cretaceous

marine sediments off the south-western Cape (McLachlan & Pieterse 1978).

Morgan (1978), also studying DSDP material, described late Cretaceous

assemblages from the Angola Basin. A brief study by Scholtz & Deacon (1982) of

sediments from kimberlite pipes in Botswana has provided evidence for the rapid

_ penetration of the ancient austral conifer forest by an early angiosperm flora

during the mid- to late Cretaceous. Sah (1967) has published a detailed

palynological study of the late Neogene sediments from Burundi, just south of the

equator. Finally, Coetzee has done pioneering work on material from numerous

short coastal sequences in the south-western Cape (Coetzee 1978a, 1981). This

work will provide the first detailed palynological evidence about the nature of

vegetation and history of vegetation changes during periods of the Neogene in

this region.

There is, however, a long history of speculative thought on the subject of

vegetation history and the evolution of certain families within the region, based

on analysis of the botanical present (Levyns 1938, 1952, 1964; Adamson 1958;

Taylor 1978; Goldblatt 1978). In the case of the more recent studies their

evidence consists mainly of present patterns of biogeography viewed against the

background of salient facts of plate tectonics. Much better use of the admittedly

patchy African fossil evidence was made by Axelrod & Raven (1978), who

combined fossil evidence, together with the two lines of evidence already

mentioned, to produce their major work on the vegetation history of Africa.

Since the fynbos vegetation of the Cape is a particularly striking component of the

vegetation of southern Africa, being accorded the status of a plant kingdom on its

own (Goldblatt 1978) and possessing a remarkable degree of endemism, and since

the early history of a number of its characteristic taxa appears to be readily

deducible from present patterns of distribution (Levyns 1964), many hypotheses

have been advanced about its origin and evolution.

It was in this context of, on the one hand, an extremely limited fossil record

and, on the other, much interest, many hypotheses and some knowledge of

biogeographical and plant taxonomic patterns relevant to an historical perspec-

tive, that a project to produce detailed palynological evidence on the composition

of vegetation in the southern African subcontinent during earlier phases of

history subsequent to the rise of the angiosperms to dominance of world floras

was undertaken. The sediments of the Arnot Pipe were known to be of late

Cretaceous to Eocene age (Estes 1977) and thus a restudy of this material was

PALYNOLOGY OF THE ARNOT PIPE 3

indicated. The importance of the Banke (Arnot) site had been emphasized by

Goldblatt (1978: 416) and such references to the sketchy evidence previously

available only increased the desirability of a thorough palynological study of the

site.

A preliminary study revealed that careful processing could produce high

pollen concentrations, that the palynomorphs were in a good state of preserva-

tion, and that the pollen assemblage was much more diverse than Kirchheimer’s

work (1934) had indicated.

As part of this project samples of lacustrine sediments were obtained from

numerous other volcanic pipes in the Namaqualand region and from further

afield, and in this paper reference is sometimes made to observations obtained in

the process of ongoing work on this material.

The aim of this study has been to systematically describe the range of forms

found in the top 20 m of polleniferous sediments (52—107 feet (c. 16-33 m) below

surface) of the Arnot Pipe and to optimize the value of these descriptions for

botanists and plant geographers by providing as many pointers to the natural

affinities of the pollen forms as was possible. It must be noted that the latter aim

was achieved almost exclusively by reference to the body of literature available to

the author. This represents an essential aspect of research, but complementary

aspects such as would be provided by the availability of a comprehensive pollen

reference collection of the floras of the subcontinent or published regional pollen

floras did not, in this study, enjoy their rightful place. The level of certainty with

which affinities are suggested is therefore variable, but this should be clear from

the text.

THE LOCATION AND GEOLOGY OF THE ARNOT PIPE

The Gamoep cluster of volcanic pipes associated with olivine-melilitite and

related rocks (Moore 1979) is located in the north-western Cape about 80 km

south of the Orange River and 100 km inland, and is centred around the hamlet

of Gamoep (Fig. 1). The cluster contains upwards of 270 pipe-like bodies

distributed with a north-north-easterly trend. It may be composed of a number of

smaller clusters (Cornelissen & Verwoerd 1975). The cluster straddles the

boundary between the Bushmanland plateau, at an elevation of around 1 200 m,

and the highly dissected escarpment area of Namaqualand. The result is that on

the plateau the pipes are typically buried under metres of sand and exposures are

poor, while in dissected country the volcanic plugs, necks or sediment-filled

depressions are exposed. The Arnot Pipe (30°22’S 18°26’E) on the farm Banke is

one of the more southerly of the pipes in the cluster and is located in a zone of

mildly dissected country between the plateau and the more highly dissected

country of Namaqualand proper. Moore (1979) states that a second distinct

cluster of pipes, associated as far as is known only with olivine-melilitite rocks, is

found between the villages of Garies and Bitterfontein, 50 km south-west of the

Gamoep cluster and closer to the coast.

4 ANNALS OF THE SOUTH AFRICAN MUSEUM

67.9m.yr

64.2m.

11.6 mr

@e® l 66,7 m.yr

oo A ® puateaxkies

GAs GOREN

\ \

‘N

1A SS

e N

Narnor, X

GARIES \

\ ;

KLIPRAND

. }

A Radiometrically dated volcanics

e Sediment filled pipes

X "Kimbernlite” pipes

. ;

BITTERFONTEIN ©

D Olivine-melilitite pipes

Fig. 1. A map of the Namaqualand—Bushmanland region of the nerth-western Cape showing the

nature and distribution of volcanic pipes in the Gamoep and Garies clusters, and the location of

dated occurrences (after Cornelissen & Verwoerd 1975; Moore 1979). Note: The majority of the

more than 270 pipes in the Gamoep cluster are not shown.

PALYNOLOGY OF THE ARNOT PIPE 5

Rogers (1911), Reuning (1931), and Cornelissen & Verwoerd (1975) have

described three categories of pipes that occur in the Gamoep cluster. Olivine-

melilitite and olivine-nepheline-melilitite pipes often form conspicuous brown,

domed hills in the dissected country. There are a limited number of occurrences

where weathered ‘kimberlite’ or ‘pseudo-kimberlite’ is exposed at the surface, but

the majority of occurrences consist of sediment and breccia-filled diatremes.

These are mostly, if not always, underlain by ‘kimberlite’ and at the present

surface display two modes of crater infilling. The minority are breccia fills in

which the sediment is disturbed by numerous large and small blocks of country

rock testifying to repeated explosive events. The majority are fills consisting of

fine-grained weathered kimberlite, carbonaceous shales and mudstones deposited

under lacustrine conditions. Doubt still remains about the exact mode or modes

of eruption of these ‘kimberlites’, which in this cluster have produced relatively

narrow pipes, some containing in the order of 300 m of bedded lacustrine

sediment. Figure 2B is a geological section through one such sediment-filled pipe,

Koppieskraal K5 (after Cornelissen & Verwoerd 1975).

The Arnot Pipe is an occurrence of the latter type. The diameter of the pipe

is 280—325 m (Reuning 1931) and it is known to contain carbonaceous mudstones

to a depth of at least 135 m (H. Jenner-Clarke, Aram Minerals, pers. comm.).

During the early 1930s a prospecting pit located towards the middle of the pipe

was sunk to a depth of 36 m to investigate the contents of the pipe (Fig. 2A). Ina

paper concerning the composition and geochemistry of the lower section of

sediments exposed in the excavation Reuning (1934) reached two conclusions of

relevance to the present work. Firstly, the sediments of the Arnot Pipe consist of

weathered ‘kimberlite’. This is an important point in suggesting the possible age

of the pipe (see section on ‘dating’). Secondly, since the lower sediments of the

pipe were composed of fine weathered and transported ‘kimberlite’ material with

an almost complete absence of derivatives from the country rock, Reuning

suggested that they were derived from the cone of a strato-type volcano that,

within the catchment area, entirely blanketed the country rock (Namaqualand

gneiss). In an earlier paper, however, Reuning (1931) had suggested another, or

complementary, reason that could explain this phenomenon as well as the lack of

heavier minerals such as ilmenite (derived from the ‘kimberlite’ itself) in the

mudstones of the pipe. A flat landscape together with the luxuriant plant growth

indicated by the rich fossiliferous nature of the sediments could have resulted in a

general low efficiency of water transport and selective deposition of finer

sediments towards the centre of the pipe. This suggestion is supported by the

work of Hawthorne (1975), who confirms that at least in the larger sedimentary

basins of the pipes found in the Botswana Kimberlite Province, coarser material is

selectively concentrated around the margins of the pipes. In the same paper

Hawthorne suggests that the actual volume of material ejected by these types of

volcanoes may have been quite small and that the cones of ejectamenta would

have been correspondingly low.

6 ANNALS OF THE SOUTH AFRICAN MUSEUM

main shaft

Be gneiss

mudstone

water table

sandstone

layers - carbonaceous mudstone

dip 30-40

dip 55°

buff coloured fine grained

“kKimberlitic’ sediments interbedded

with carbonaceous mudstone.

rich in macrofossils, frogs. leaves

and branches

fewer frogs, but richer accumulation

of leaves

80 ft

arkose and grit

carbonaceous mudstone

conglomerate

tuffaceous

“kimberlite”

"kimberlite’with

blocks of Country

rock

Fig. 2. A. A stratigraphic section of the known sequence of the Arnot Pipe. The upper 33 m

(107 feet) (maximum depth of the 1929 excavation) at the main shaft and the side excavations are

shown after Reuning (1931). (Reuning’s measurements are given in feet.) The provenance of

samples examined in this study is indicated. The carbonaceous mudstones were recorded to a

depth of 135m in drilling done by H. Jenner-Clarke (pers. comm.) during the 1960s.

B. A stratigraphic section based on the logs of two cores of the Koppieskraal K5 pipe. This can be

taken as a representative reconstruction of the stratigraphy of sediment-filled pipes in the

Gamoep cluster (after Cornelissen & Verwoerd 1975).

PALYNOLOGY OF THE ARNOT PIPE 7

A final point in respect of the origin of the fine sediments of the pipe and the

lack of coarser sediments needs to be made. In Reuning’s (1931, 1934) arguments

he assumed that the substrate in the area at the time of deposition would have

been the country rock, Namaqualand gneisses. However, inclusions of Dwyka

shales have been observed in pipes (Moore 1979: 9) and in the present study an

odd palynomorph specimen of Permian—Triassic provenance was observed. This

indicates that at the time when sediments were being deposited in the crater lake

some Dwyka cover was still present in the area.

From this survey of what is known of the geology of the pipes the following

picture emerges of the local environment during the time of eruption and

sedimentation of the pipes. A great many strato-type ‘kimberlite’ volcanoes

erupted during a relatively short time in a small area. (Using an estimate of a total

of at least 350 pipes in a region of 8 400 km? and the present span of radiometric

dates of 7 Ma, the following calculations can be made: The average density of

pipes is one per 24 km’, though they may be concentrated in subclusters with

densities of about one per 4 km’; average time between eruptions 17 000 years.)

Many small crater lakes could have been synchronously present in the region

although, taking into account the ease with which the ejectamenta could have

weathered, each crater may have had a relatively short life of sediment capture

before infilling was completed.

Using the range of estimates of sedimentation rates for alluvial fan sediments

given by Hooke (1968) and Beaty (1970) it would require between 300 000 and

4 000 000 years to accumulate the possible depth of sediment present in the Arnot

Pipe (135-300 m). The rate of sedimentation, however, would decrease as the

cone height and supply of tefra was reduced.

The basins would presumably have remained as swamp-like features for a

longer period of time and, due to compaction or shrinkage of the initial

sedimentary mass, may have retained a minor ability to capture sediments. The

steeply inward-dipping strata often observed in the pipes (see Fig. 2) is evidence

that this may have occurred. The small drainage basins feeding the lakes would

have been largely unrelated to the developed drainage patterns of the region.

The general landscape was probably dominated by the small emergent cones

of the ‘kimberlite’, strato-type volcanoes in various stages of erosion and, for the

rest, there is little reason to suppose that a relatively flat, mature landscape did

not exist (Mabbutt 1955). Although in the present the peaks of the nearby

Kamiesberge rise some 150-200 m above the plateau, most of the present relief

of Namaqualand is probably the result of more recent incision.

PREVIOUS STUDIES OF FOSSIL MATERIAL

FROM THE ARNOT PIPE

During the 1930s, Reuning and later Boonstra selected samples of material

from the dump of the main excavation. Boonstra (unpublished notes in the South

African Museum) roughly indicated the stratigraphic provenance of a series of

8 ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 1

Material from the 1929 excavation of the Arnot Pipe collected and provenanced by E. Reuning

and L. D. Boonstra and analysed in this study.

Provenance of sample Description of the material

52-58 feet (16-18 m) Bituminous, carbonaceous mudstone.

Unprovenanced Bituminous, carbonaceous mudstone.

65-70 feet (20-21 m) Buff-coloured finely laminated mudstone.

70-90 feet (21-27 m) Buff to grey-coloured mudstone.

90-100 feet (27-30 m) Buff-coloured mudstone with fine silty laminations.

100-107 feet (30-33 m) Buff-coloured mudstone.

100-107 feet (30-33 m) Brown-coloured mudstone with large flecks of organic material.

five small samples (see Fig. 2A and Table 1). A considerable amount of material

was deposited at the South African Museum and the following categories of fossil

material were later studied by various researchers: leaves (Rennie 1931), wood

(Adamson 1931), frogs (Haughton 1931), and palynomorphs (Kirchheimer 1934).

The fossil leaf collection consisted of 70 fragments of dicotyledonous leaves

(and a single fern frond) and Rennie recognized at least twelve different types. In

his short paper only six forms are illustrated by rough line-drawings. The most

common form was described as strap-shaped with serrate margins, and although

Rennie did not do so, it could be described as sclerophyllous. This form was

tentatively compared to the leaves of Myrica, the comparison to some extent

being based on identifications contained in the work of Berry (1925) on Upper

Cretaceous leaf assemblages from North America (Rennie 1931: 252). Rennie’s

tentative comparison is rendered less likely by Chourey’s (1974: 131, 145)

thorough criticism of Berry’s work. Chourey noted that in the early years of study

of leaf fossils of late Cretaceous—Tertiary age, inadequate identification criteria

were employed and many identifications made then can no longer be accepted. In ~

particular many fossils were identified either as myricaceous or proteaceous;

affinity with the genus Banksia of the Proteaceae was regularly suggested. In fact

this tentative alternative identification was made by Berry in the very paper cited

by Rennie. It should be noted that, while rejecting most of the Myrica identifi-

cations made on late Cretaceous—early Tertiary material from North America and

Europe, Chourey (1974) suggests that the phenomenon of a world-wide occurrence

and prominence of this form type is significant and worthy of further study. The

‘myricaceous’ leaves from Arnot are part of this world-wide phenomenon.

It was not possible to obtain adequate descriptions of the remaining eleven

leaf types to allow for identification and Rennie merely noted their general

‘mesophytic’ habit.

Adamson (1931) studied silicified wood samples from opalized sections of the

superficial sandstone layers found close to the contact between the pipe fill and

the gneiss. He identified the fossil wood as that of Ficus cordata, which grows in

the area at present. Kirchheimer (1934: 47) quotes Reuning’s statement that these

sandstone layers may be of a substantially younger age than the underlying clays.

PALYNOLOGY OF THE ARNOT PIPE 9

Haughton (1931), in the most detailed of these early studies, described a

single new species of Pipidae, Eoxenopoides reuningii and, in terms of relatively

conservative pipid evolution and lacunae in their fossil record, could only assign a

Cretaceous to early Tertiary age to the form. Estes (1977) agreed with

Haughton’s systematic description and on morphological grounds also upheld

Haughton’s age bracketing. However, on extraneous grounds he favoured an

Oligocene age for the sediments (Estes 1977: 51).

Kirchheimer (1934) cursorily described six palynomorph types including two

disaccate and four triaperturate angiospermous forms. He also mentioned that

spores and inaperturate grains had been observed. He obtained his best results

from thin sectioning of opal concretions and obviously encountered difficulties in

processing the carbonaceous clays of the lower layers available to him.

DATING

The best biostratigraphic evidence on which to base an age estimate for the

sediments of the Arnot Pipe has been produced by the present study and is

discussed in a later section in the context of the radiometric evidence mentioned

below. The suggestion by Axelrod & Raven (1978), based on an extremely

tenuous comparison with the North American and Mediterranean plant macro-

fossil record, that the occurrence of a sclerophyllous leaf type indicates a late

Eocene to Miocene age, can be regarded as insubstantial. The arguments

advanced by Reuning (1931) and Haughton (1931) on sedimentological and

geomorphological grounds for a late Cretaceous age were probably never

intended to be more than speculative.

The most positive dating evidence relevant to suggesting a possible age for

the Arnot sediments consists of a number of radiometric determinations on

material from pipes in the Gamoep cluster (Davis 1977). The dates and location

of pipes are given in Figure 1. All the dates were obtained by the *°U/*"°Pb

method applied to zircon inclusions in kimberlitic material. Until these dates were

available, the single K/Ar date of 38,5 Ma from an olivine-melilitite pipe on the

farm Dikdoorn in the Garies—Bitterfontein cluster was the most pertinent

radiometric determination relevant to the possible age of the Gamoep cluster.

This was also the only date available when the dating of the Arnot sediments was

last discussed (Estes 1977).

In view of the range of dates presently available, the evidence is strongly in

favour of accepting a 60-70 Ma age bracket for volcanic activity in the Gamoep

cluster and for the Arnot Pipe. Firstly, all three pipes dated in the Gamoep cluster

are apparently ‘kimberlitic’, as is the Arnot Pipe. Secondly, the five dates

available are relatively tightly grouped within a 7 Ma time span and the Arnot

Pipe is within 30 km of the dated pipes. No olivine-melilitite pipes from the area

have yet been dated.

Two K/Ar dates from olivine-melilitite pipes in the small cluster south of

Garies are available and are younger or considerably younger than the dates for

10 ANNALS OF THE SOUTH AFRICAN MUSEUM

the main Gamoep cluster, i.e. 54,1 and 38,5 Ma. It must be pointed out that the

younger date here and the other two Oligocene dates shown in Figure 1 have in

fact not been fully published (Kréner 1973) and should be treated with caution. In

contrast to the use of the *°U/*%°Pb method to date the pipes in the Gamoep

cluster, the K/Ar method was used in the case of the Garies—Bitterfontein pipes.

Excluding possible problems involved in the dating techniques, the spread of

dates from the Garies—Bitterfontein pipes as well as from olivine-melilitite pipes

wider afield (see Fig. 1) may indicate that the time span of volcanic activity either

in this small cluster or throughout the distribution of olivine-melilitites was much

greater than that of the volcanic activity associated with the ‘kimberlite’ pipes of

the Gamoep cluster. Moore (1979) (see below) has suggested at least two

mechanisms that could explain the phasing of volcanic activity resulting in

extrusion of lighter ‘kimberlite’ earlier than that of more dense olivine-melilitite.

He (1979: 136) also discusses some disparities in results obtained in the

radiometric dating of kimberlites, olivine-melilitites and related rocks, and

suggests that many determinations are possibly questionable.

Moore (1973, 1979) develops hypotheses that might explain:

(a) the possible younger ages of the olivine-melilitite versus the ‘kimberlite’

volcanics of the region;

(6) the possible pattern of younger dates occurring closer to the coast; and

oxide content of volcanics from the olivine-melilitites of the Garies—

Bitterfontein cluster through those of the Gamoep cluster to the ‘kimberlites’

of the latter region.

These hypotheses involve the late Cretaceous—Oligocene epeirogenic uplift

along the warp axis of the western escarpment as a primary cause of volcanic

activity. Furthermore, either the fractionation of the parent magma in the stress

zone beneath the warp and a resultant enrichment of a less dense portion with

volatiles could lead to a first phase of volcanic activity in which ‘kimberlite’ was

extruded, or the progressive thickening of the craton towards the interior of the

continent might cause extrusion of magmas from different depths and thus

explain the coast—interior gradient of geochemical attributes and perhaps dates.

So much for the possible dating of the volcanics that provides a maximum age

for the lacustrine deposits. As suggested it is likely that infilling of the craters

would have proceeded relatively rapidly, or would at least at first have been

rapid. The estimates obtained from sedimentological work in similar environ-

ments suggest that a maximum of 4 Ma could have been required to accumulate

the depth of deposit that may be present in the Arnot Pipe.

MATERIAL AND METHODS

As already stated, the material curated in the South African Museum was

collected by E. Reuning and L. D. Boonstra during the 1930s. Even the five small

samples whose stratigraphic provenance was indicated by Boonstra (see Fig. 2A

PALYNOLOGY OF THE ARNOT PIPE 1]

and Table 1) were probably collected from the dump (which was presumably to

some extent systematically organized) after the excavation had been closed. This

is inferred from the fact that this small series includes samples from the lowest

levels reached in the excavation, yet the work had ceased at this level (35 m) due

to the instreaming of water, and the pit would presumably have filled with water

to the level of the water-table 20 m higher up. The coherent pattern of the pollen

diagram (Fig. 3) does to some extent suggest that the provenancing of the samples

is correct.

The seven samples studied are listed and described in Table 1, in the same

sequence as their pollen spectra appear in the pollen diagram.

The bulk of the collection was unprovenanced, but since this material often

contains macroscopic fossil material, it is unlikely to have come from the upper

third of the excavation. Reuning (1931) noted that preservation of fossil material

above the level of the water-table was poor. On the other hand, in view of the

pollen counts (Fig. 3), it seems unlikely that the bulk of the samples could have

come from below the 18 m (60 foot) level. The combined count of two counts on

the unprovenanced material is placed below those of Boonstra’s 52-58 foot

(16-18 m) samples on the pollen diagram and are virtually indistinguishable from

the latter.

The unprovenanced material is of two sedimentological types—a dark,

carbonaceous mudstone and a buff-coloured mudstone. The pollen spectra from

the two types are similar and unfortunately, since the best-preserved and richest

concentrations were obtained from this material, many holotypes designated in

this study are located in preparations from it.

It appeared that two sediment types also occurred between 30 and 33 m

(100-107 feet), but no indication was given about their stratigraphic relationship.

In Table 1 and Figure 3, the spectrum from the buff-coloured mudstone 1s

arbitrarily placed above that of the dark, carbonaceous mudstone.

Kirchheimer used a single-stage process, cold hydrofluoric acid digestion, to

concentrate palynomorphs. The stratigraphic provenancing of his samples was

uncertain. In the present study a six-stage process was employed, consisting of the

following steps:

1. Crushing and dispersal of the sample in a 0,3M solution of tetrasodium

pyrophosphate, followed by numerous short centrifuges and rinses in deionized

water to wash out the fine clay fraction. If necessary, additional applications of

tetrasodium pyrophosphate solution were used. This treatment is an adaptation

of the process described by Bates et al. (1978) but was independently suggested by

a soil scientist, J. J. N. Lambrechts of the Department of Agriculture, University

of Stellenbosch.

2. Standard zinc chloride heavy-liquid flotation to separate organic and

larger-sized inorganic fractions.

3. In the treatment of a few of the more heavily carbonaceous samples it was

necessary to cause some oxidation using either 30 per cent hydrogen peroxide

solution or nitric acid.

[2 ANNALS OF THE SOUTH AFRICAN MUSEUM

eS 3s ¢ & SSeS a ee Ss

, 1 = © es ) w o co a 8 4! > ~

as" 38 Ss 2 8 4 via: eo en tee PROVENANCE OF SAMPLE

— ~ . - —_- = oe o Qo mm [.s}

PS = — a — a = ] e - =< e@

= = = = =

oO = = oO

= =

STEREISPORITES TRIORITES OPERCULATUS

OTHER SPORES

| J DICOLPOPOLLIS

TRICOLPITES RETICULATUS

CLAVATIPOLLENITES

TRIORITES SPHERICUS

PROPYLIPOLLIS

ERICIPITES

PODOCARPIDITES

NILFOROIA

“ LE he 8 ARAUCARIACITES

™m

TRICOLPOROPOLLNITES ARNOTIENSIS

[=]

E a R i R ry Wm TRICOLPOROPOLLENITES BRINKIAE

S233 S & S EB poruen sun GT OW TM ™ rerpororerravires spuerrcus

Fig. 3. Pollen diagram showing relative abundance of those taxa that constitute greater than

1 per cent of the grains present at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 1

4. Standard acetolysis process.

5. A two or three-minute rinse in hydrofluoric acid to destroy any remaining

silica particles.

6. A rinse in warm 10 per cent hydrochloric acid.

After final washing with deionized water the concentrates were suspended in

a 50 per cent glycerol solution for light microscopy or left in deionized water for

mounting on SEM stubs. Permanent slides were made with glycerine jelly and

sealed with nail varnish.

The full series of slides with holotypes and paratypes ringed and documented

is deposited in the South African Museum. A register of South African Museum

catalogue numbers for these holotypes and paratypes is given in the species list

(e.g. SAM-K6155). A duplicate set of slides and the sealed phials containing the

remaining concentrates is deposited in the Department of Archaeology,

University of Stellenbosch. Counting of samples and photomicroscopy was done

on a Wild M11, and SEM work was done on a JEOL JSM-35 housed in the

Department of Physics, University of Stellenbosch.

PALYNOLOGY

Kemp & Harris (1977: 5) provide a recent review of the problems

encountered in the systematic palynology of early Tertiary material. They state

that “Tertiary palynology even more than the palynology of older sedimentary

rocks, has suffered from a marked ambiguity of approach’. This ambiguity of

approach arises in that three different approaches to nomenclature have been

applied. Authors have variously assigned fossil palynomorphs to extant genera,

used a name that suggests affinity to an extant genus (e.g. Araucariacites for forms

resembling pollen of the genus Araucaria), or applied an artificial name based on

morphological criteria alone (e.g. Triorites or Monocolpopollenites).

In addition, a certain regionality of nomenclature, reflecting the isolation in

which early palynological work was done, is inherited by present-day palynol-

ogists (depending on what literature they are exposed to). This problem can be

compounded by the difficulties entailed in keeping up to date with more recent

work published in an array of journals.

Most palynologists working on early Tertiary material have elected, firstly, to

continue to use the binomial system of nomenclature together with the standard

botanical rules of typification, priority, etc. This promotes stability and some

measure of uniformity in the use and creation of names for fossil palynomorphs,

and is the system adhered to in the present study. However, it does not

discourage the proliferations of names and the growth of ‘portmanteau’ genera.

Secondly, in establishing new genera palynologists have favoured the artificial

system of nomenclature, which promotes the utilization of palynology as a strati-

graphic tool (Sah 1967: 6), and this practice is also followed in the present work.

In this study the descriptive and stratigraphic palynological literature from

Australia, India and tropical Africa has been most often consulted. As could be

14 ANNALS OF THE SOUTH AFRICAN MUSEUM

expected, in view of its isolation from these areas, the palynomorph assemblage

from Banke appeared unusual and many new specific names have resulted. Both

in the occurrence of a number of unique forms and in the general composition of

the assemblage, the already developed distinctiveness of the flora of the

subcontinent is apparent.

A new specific or generic name was not proposed unless the form concerned

was reasonably common in the Arnot samples. A large number of forms were

present at low frequencies and in these cases, if at least three specimens in good

state of preservation were observed, the forms were described and illustrated,

placed if possible in a genus, and their affinities suggested. This was deemed to be

worthwhile in terms of the aims of the study and the unique nature of these

observations at present. Other forms will no doubt be systematically described as

work on the pollen and spore assemblages from ‘“kimberlite’ pipe occurrences in

the northern Cape and Botswana is extended. Sequences are already known in

which elements rare at Arnot are common or even dominant. One form, referred

to informally as Fenestriorites, is not described in the present paper as it is being

described elsewhere. The form promises to be an especially important marker

species in the regional sequence. Some importance is therefore placed on its

occurrences in the lowest levels as yet sampled at Arnot (where it is extremely

rare) and this is discussed in the section on interpretation of the present

palynological evidence.

SPECIES LIST AND REGISTER OF TYPE SPECIMENS

Spores

Trilete spores

Stereisporites sp.

Cyathidites australis Couper, 1953

Planisporites sp.

Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968

Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973

Foraminisporis sp.

Herkosporites elliottii Stover, in Stover & Partridge, 1973

Camarazonosporites bankiensis sp. nov.

Holotype SAM-K6155

Polypodiaceoisporites sp.

Trilites sp.

Monolete spores

Microfoveolatosporis fromensis (Cookson) Harris, 1965

Cicatricososporites sp.

Alete spores

Reticulatasporites grandis sp. nov.

Holotype SAM-K6156; paratypes SAM-—K6157, K6158

PALEY NOLOGY OFSHHE ARNOT PIPE

Spores not assigned to genus

Forma A

Forma B

Forma C

Forma D

Pollen of Coniferae

Inaperturate pollen

Araucariacites australis Cookson ex Couper, 1953

Araucariacites sp.

Monosaccate pollen

Zonalapollenites sp. A

Zonalapollenites sp. B

Disaccate pollen

Lygistepollenites sp.

Podocarpidites sp.

Podocarpidites kamiesbergensis sp. nov.

Holotype SAM-K6159; paratypes SAM—K6160, K6161, K6162

Podocarpidites riembreekensis sp. nov.

Holotype SAM-K6163; paratypes SAM-—K6164, K6165, K6166

Pollen of Angiospermae

Monoaperturate pollen

—Monocolpate pollen

Arecipites plectilimuratus Chmura, 1973

Arecipites sp. A

Arecipites sp. B

Liliacidites sp. A

Liliacidites sp. B

Clavatipollenites sp. A

Clavatipollenites sp. B

Clavatipollenites sp. C

Monocolpopollenites sp. A

Monocolpopollenites sp. B

—Monoporate pollen

Milfordia hypolaenoides Erdtman, 1960

Milfordia sp.

—Dicolpate pollen

Dicolpopollis sp.

—Monocolpate pollen not assigned to genus

Forma E

16 ANNALS OF THE SOUTH AFRICAN MUSEUM

Triaperturate pollen

—Triporate pollen

Triorites operculatus sp. nov.

Holotype SAM-—K6167; paratypes SAM—K6168, K6169, K6170

Triorites sphericus sp. nov.

Holotype SAM-K6171; paratypes SAM—K6172, K6173

Triorites harrisii Couper, 1960

Triporopollenites namaquensis sp. nov.

Holotype SAM-—K6175; paratypes SAM-—K6176, K6177, K6178

Proteacidites sp. A

Proteacidites sp. B

Propylipollis meyeri sp. nov.

Holotype SAM-K6179; paratypes SAM-—K6180, K6181, K6204

Propylipollis sp.

Fenestriorites sp. (not described in this paper)

—Tricolpate pollen

Tricolpites reticulatus Cookson, 1947

Tricolpites sp. A

Tricolpites sp. B

Tricolpites sp. C

Tricolpites sp. D

Crototricolpites densus Salard-Cheboldaeff, 1978

Spinitricolpites jennerclarkei gen. et sp. nov.

Holotype SAM-K6182; paratypes SAM—K6183, K6184, K6185

—Tricolporate pollen

Tricolporopollenites grandis sp. nov.

Holotype SAM-K6186; paratypes SAM-K6187, K6188

Tricolporopollenites arnotiensis sp. nov.

Holotype SAM-K6189; paratypes SAM—K6190, K6191

Tricolporopollenites brinkiae sp. nov.

Holotype SAM-K6193; paratypes SAM—K6194, K6195

Tricolporopollenites coetzeeae sp. nov.

Holotype SAM-K6196; paratypes SAM-K6192, K6174

Tricolporopollenites sp. A

Tricolporopollenites sp. B

Tricolporopollenites spp. C & D

Pollen with more than three apertures

Retistephanocolpites sp.

Grootipollis reuningli sp. nov.

Holotype SAM-K6197; paratypes SAM-—K6202, K6203

Ulmipollenites sp.

PALYNOLOGY OF THE ARNOT PIPE 17

Inaperturate pollen

Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979

Pollen found in obligate tetrads

Ericipites sp. A

Ericipites sp. B

Dicotetradites sp.

Triporotetradites sphericus sp. nov.

Holotype SAM-—K6198; paratypes SAM-—K6199, K6200, K6201

Dicotyledononous pollens not assigned to genus

Forma F

Forma G

Forma H

Forma I

DESCRIPTIONS

Spores

‘The identification of Tertiary spores is more problematical than that of fossil

pollen grains. This is partly because they are less well known but mainly because,

as Knox (1935) and Selling (1946) have shown, a particular type is not necessarily

restricted to a single genus or even family and considerable variation often exists

within a genus’— Cookson (1947: 135).

Trilete spores

Genus Stereisporites Pflug, in Thomson & Pflug, 1953

For a discussion of this genus see Dettmann (1963: 25).

Stereisporites sp.

Fig. 4A—D

Compare

Sphagnum antiquasporites Wilson & Webster, 1946: 273 (fig. 2).

Triletes australis Cookson, 1947: 136, pl. 15 (figs 58-59).

Sphagnites australis (Cookson) Balme, 1957: 15, pl. 1 (figs 1-3).

Stereisporites antiquasporites (Wilson & Webster) Dettmann, 1963: 25,

figs 20-21. Harris 1974: 79, pl. 24 (fig. 20).

Description

Microspore trilete, biconvex, amb subtriangular to subspherical with convex

sides and broadly rounded angles. Laesurae straight and simple, length one-half

spore radius. The exine is uniformly thick and the distal and probably the

proximal surfaces are covered by very low angular rugulae. Equatorial diameter

24-35 pw, exine 1-2 wm.

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. A-D. Stereisporites sp. EE. Cyathidites australis. _F—G. Planisporites sp.

H. Foveotriletes margaritae. \—J. Foveotriletes lacunosus. K—L. Foraminisporis sp.

PALYNOLOGY OF THE ARNOT PIPE 19

Remarks

The slight thickenings in the radial regions at the equator and the low, distal

polar thickening, circular in outline, mentioned by Dettman (1963: 25) were not

observed, nor do they appear in either the descriptions or photomicrographs of

Wilson & Webster (1946), Cookson (1947), Balme (1957) or Harris (1974). The

differences between the present form and the species Stereisporites antiqua-

sporites (Wilson & Webster) Dettmann, 1963, are therefore slight, but sufficient

to prevent identification.

Affinity

The genus Sphagnites Cookson, 1953, was established to include fossil spores

resembling those of the peat-moss family Sphagnaceae. (See discussion of

Stereisporites in Boros & Jarai-Komlddi 1975: 9.)

Distribution

The genus is known from Jurassic to Tertiary sediments and is sometimes

common in Australian late Cretaceous sediments, particularly in highly carbon-

aceous samples (Dettmann 1963). Stereisporites sp. is the most common spore at

Banke. Due to the long time range shown by the genus Stereisporites, it may have

little stratigraphic value, but Balme (1957: 15) remarks that it is rarely seen in

marine and transitional sediments and may therefore be important in facies

studies. A Sphagnites form is illustrated but not described from the Knysna

lignites (Neogene?) by Thiergart et al. (1963, table 2, fig. 6) and was also recorded

in sediments of middle to late Cretaceous age off the south-western Cape

(McLachlan & Pieterse 1978). A survey of the literature suggests that Sphagnites

forms are not known from late Cretaceous or Tertiary sediments of tropical

Africa. The modern family is strongly circumboreal in distribution, although one

form is cosmopolitan and other forms occur in the Southern Hemisphere, mainly

in New Zealand (Boros & Jarai-Komlddi 1975). A few Sphagnum species are

found in southern Africa and are common although ‘confined to shaded mountain

seeps, streambanks or swampy areas’ (Magill 1981: 23).

Genus Cyathidites Couper, 1953

See discussion in Dettmann (1963: 22).

Cyathidites australis Couper, 1953

Fig. 4E

Cyathidites australis Couper, 1953: 27, pl. 2 (fig. 11).

Description

The spores are trilete. The laesurae are ciearly defined (about two-thirds of

the radius of the spore), narrow and straight. The ends of the laesurae may

20 ANNALS OF THE SOUTH AFRICAN MUSEUM

terminate in a short bifurcation. The spores are triangular with rounded apices

and the sides are mostly concave in polar view. The exine is thin and psilate, and

both proximal and distal surfaces are convexly curved. Equatorial diameter

46-60 pw, exine 1 p.

Affinity

Couper (1953: 27), having compared the present form to the spores of the

extant fern Thyrsopteris elegans, quoted Copeland (1947: 48) on the latter

species: ‘It may well be a relict from the time when Dicksonia and Cyathea had a

common ancestor.’ On the basis of the available evidence Couper suggested that

Cyathidites australis may be the spore of a tree-fern.

Distribution

Cyathidites australis is widely distributed in the Mesozoic and Tertiary of the

Northern Hemisphere and Australia, and is often abundant. It is a rare

component of the Banke spore flora. Similar forms are present in the late

Cretaceous (McLachlan & Pieterse 1978, pl. 1 (figs 1-2)) and Neogene (?)

(Thiergart et al. 1963, pl. 2 (figs 12, 14—15)) of the southern African subcontinent.

Genus Planisporites Knox, 1950

Planisporites sp.

Fig. 4F-—G

Description

The spore is trilete and its amb is subtriangular to deltoid with rounded

corners. The exine is relatively thick. The distal face and equatorial regions are

decorated with microconi, which are mostly solitary, but may also be arranged in

rows. The bases of adjacent coni may be linked by fine ridges. The proximal face

is psilate and the laesurae are long, thin and simple and almost reach the equator.

The equatorial diameter of the illustrated specimen is 33 w and the exine is 2 wu

thick.

Affinity

There is no information regarding possible affinities of Planisporites sp.

Distribution

Planisporites sp. is rare at Arnot. There is no further information on the

distribution of the genus.

PALYNOLOGY OF THE ARNOT PIPE 2A

Genus Foveotriletes Potonié, 1956

Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968

Fig 4H

Triletes margaritae van der Hammen, 1954: 102, pl. 17.

Foveotriletes margaritae (van der Hammen) Germeraad et al., 1968: 286, pl. 1 (figs 1-2).

Description

The spore is trilete with a roundly triangular amb and is circular to biconvex

in lateral view. The laesurae are straight with finely serrate margins and are one-

half the spore radius. The exine is relatively thin and the whole surface is densely

covered by scrobuli; approximately sixty scrobuli per 100 «*. Equatorial diameter

is 55 w; laesurae 15-18 w; exine 1-2 wp.

Affinity

Spores that resemble F. margaritae are produced by members of the

Ophioglossaceae, especially the genera Botrychium and Ophioglossum (Salard-

Cheboldaeff 1981).

Distribution

Foveotriletes margaritae is recorded from sediments of Palaeocene age in

tropical Africa and South America, where it becomes extinct during the lower

Eocene (Germeraad ef al. 1968). It is rare at Arnot.

Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973

Fig. 4I-J

Foveotriletes lacunosus Partridge, in Stover & Partridge, 1973: 248, pl. 14 (fig. 6).

Description

The spore is trilete and the amb is rounded triangular to subcircular. The

distal surface is convex and the proximal is pyramidal. The laesurae are two-thirds

to three-quarters spore radius, irregular, not straight, and have thin, steep lips.

The proximal surface is psilate, the distal surface is covered with poorly

delimited, shallow foveola that almost encroach on to the proximal face. The

exine is relatively thick, approximately 2 uw, and its inner surface appears to

follow the undulations of its outer surface. Equatorial diameter 35-37 wp.

Affinity

There is no information on the affinity of F. lacunosus.

Distribution

Foveotriletes lacunosus is known from the Oligocene to Miocene in

Australia; it is rare at Arnot.

22 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Foraminisporis Krutzsch, 1959

See discussion in Dettmann (1963: 71).

Foraminisporis sp.

Fig. 4K—L

Compare

Foraminisporis dailyi Dettmann, 1963: 72, pl. 14 (figs 15-18).

Description

The spore is trilete. The amb is rounded triangular to subcircular with a

notch in what is probably a narrow sculptured cingulum (see Dettman 1963: 71)

where the laesurae meet the equator. The outline of the grain is irregular. The

laesurae are straight and placed on sculptured ridges running the length of the

radial areas. The distal face is covered by verrucate to spinulate structures whose

bases sometimes coalesce. The proximal face is distinctly less verrucate with

occasional foveola. Equatorial diameter 45-50 p, cingulum 3 wp.

Affinity

The affinity of the genus Foraminisporis is perhaps with the bryophyte family

Anthocerotaceae (Dettmann 1963: 71).

Distribution

Foraminisporis 1s world-wide in the Cretaceous; F. dailyi is present in late

_ Cretaceous sediments off the south-western Cape (McLachlan & Pieterse 1978).

Foraminisporis sp. was common at Arnot.

Genus Herkosporites Stover, in Stover & Partridge, 1973

See discussion in Stover & Partridge (1973: 248).

Herkosporites elliottii Stover, in Stover & Partridge, 1973

Fig. 5A—B

Herkosporites elliottii Stover, in Stover & Partridge, 1973: 248, pl. 13 (fig. 7).

Description

The spore is trilete, the amb roundly triangular and the laesurae extend

almost to the equatorial margins. The radial area immediately bordering the lips

of the laesurae appears smooth, but in the remainder of the radial region

regularly spaced, thin structures (? folds) arranged at right angles to the laesurae

are present. The proximal interradial area is psilate. The laesurae are narrow with

thin raised lips. The distal surface is spinate; spines are of a uniform height 3-4 yu

with abruptly broadening bases, which sometimes coalesce in fine ridges; space

between spines 1,5—2,5 uw, exine 1 w, equatorial diameter 40-45 w.

PALYNOLOGY OF THE ARNOT PIPE 23

Affinity

There is no direct information on the affinities of Herkosporites. Dettmann

(1963: 36) quotes Cookson & Dettmann (1958) on a comparison between the

morphologically related genus Ceratosporites and certain members of the extant

genus Selaginella.

9046

Fig. 5. A-B. Herkosporites elliottii. C-D. Camarazonosporites bankiensis sp. nov.

E-F. Polypodiaceoisporites sp.

24 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Herkosporites elliottii is distributed in the Palaeocene to Miocene in

Australia; it was rare at Arnot.

Genus Camarazonosporites Pant ex Potonié, 1956

Camarazonosporites bankiensis sp. nov.

Fig. SC-D

Etymology

This species is named after the farm Banke, near Platbakkies, Namaqualand,

and the site name.

Description

The spores are trilete and cingulate with a convexly triangular amb and

rounded apices; biconvex in lateral view. The proximal face is subpsilate,

probably finely granulate. The distal face is covered by a fine hamulate

sculpturing, which is completely lost in the radial equatorial region, following the

characteristic trend for the reduction of the exine of this region in the genus

Camarazonosporites. The narrow laesurae extend to the equator and are

bordered by thin, steep membraneous folds, which thus form irregular lips. The

lips may overfold the laesurae. The cingulum (equatorial crassitude) is 6-7 pu

wide in the interradial regions, and narrows to 1-1,5 w in the radial region, giving

the grain a rounded triangular aspect in plan view. Equatorial diameter is

59-63 p.

Remarks

Camarazonosporites bankiensis is quite common in the Banke material and

can therefore serve as a type population. The spores are twice the size of the type

species of the genus, and much larger than any other known species of

Camarazonosporites. It is also larger than most species in morphologically related

genera such as Coronatispora, Sestrosporites and Camarazonotriletes.

Affinity

The affinities of C. bankiensis are not known. It has a general similarity to

the spores of some members of the Lycopodiaceae.

Distribution

Camarazonosporites bankiensis is common at Arnot. The genus is known

from the late Cretaceous. An apparently similar form, labelled Lycopodium-

type, is present in the Knysna lignites (Neogene?) (Thiergart ef al. 1963, pl. 2 (figs

1—3)) but no description is provided.

PAL Y NOLOGY OPMHE ARNOT PIPE 25

Genus Polypodiaceoisporites Potonié, 1951 ex Potonié, 1956

Polypodiaceoisporites sp.

Fig. SE-F

Description

A trilete, cingulate spore. The amb is triangular with broadly rounded

apices. The laesurae are straight and reach the cingulum without extending into

it. A concavely triangular area in the central radial area of the proximal face is

markedly depressed below a surrounding ridge, which is decorated with robust

rugulate structures. This sculpturing becomes reduced as the laesurae are

approached within the centrally depressed area. The rugulate sculpturing on the

distal face is formed by much flatter and broader structures. The cingulum is

smooth and of variable width. Equatorial diameter 80 w, cingulum 6 w wide.

Affinity

The generic designation suggests affinity to the Polypodiaceae, but the spores

of the genus Pteris of the Pteridaceae (Muller 1968) also resemble this form.

Distribution

The genus Polypodiaceoisporites is known from the Tertiary and Cretaceous

of both the Southern and Northern hemispheres. Polypodiaceoisporites sp. is rare

at Arnot.

Genus Trilites Cookson ex Couper, 1953

Trilites sp.

Fig. 6A-B

Compare

Trilites ohaiensis Couper, 1953 in Couper, 1960: 41, pl. 2 (figs 7-8).

Latrobosporites crassus Harris, 1965: 81, pl. 25 (figs 8-9).

Description

The spore is large and trilete with a relatively thick psilate exine. It is

uniformly covered by a thin outer membrane, which closely adheres to the exine

and is thrown up into low folds and wrinkles to form a dense hamulate pattern.

The laesurae are long and thin and outlined by folds in the perinous membrane.

The laesurae reach or almost reach the equator. It appears that the perinous

membrane withdraws from a small area around the apices of the amb where the

laesurae meet at the equator. This suggests that the grain is limbate. The amb of

the grain is that of a broadly rounded triangle. Equatorial diameter 60 w.

Fig. 6.

ANNALS OF THE SOUTH AFRICAN MUSEUM

i ‘ 1 n 40 se

A-B. Trilites sp. C. Microfoveolatosporis fromensis. D. Cicatricoso-

sporites sp. E-F. Reticulatasporites grandis sp. nov.

PALYNOLOGY OF THE ARNOT PIPE 27

Remarks

Despite Harris’s (1965: 81) comment that no forms resembling the new genus

and species Latrobosporites crassus Harris, 1965, were then known from Austral-

asia, there seems to be some measure of resemblance between TJrilites ohaiensis

Couper, 1953 and L. crassus Harris, 1965. There is a resemblance between these

two species and the present form, Trilites sp., although the amb of the latter is

more triangular and neither of the former two species are described as perinate.

Trilites ohaiensis was originally described (Couper 1953: 30) as having a

verrucate—granular sculpture, but this was later revised (Couper 1960: 41) and the

grains described as having a rugulate—vermiculate sculpture. Harris (1965)

described the sculpture of L. crassus as consisting of low interlocking rugulae and

lumina of similar size and shape.

Affinity

Harris (1965) suggested that L. crassus had affinity to the extant Selaginella

cathedrifolia-group as defined by Knox (1950).

Distribution

Trilites sp. is rare at Arnot. Trilites ohaiensis is rare in New Zealand late

Cretaceous sediments (Couper 1960), and L. crassus is common in Palaeocene

sediments from south-western Australia (Harris 1965).

Monolete spores

Genus Microfoveolatosporis Krutzsch, 1959

Microfoveolatosporis fromensis (Cookson) Harris, 1965

Fig. 6C

Schizaea fromensis Cookson, 1956: 43, pl. 8 (fig. 3).

Microfoveolatosporis fromensis (Cookson) Harris, 1965: 84, pl. 24 (fig. 7).

Description

The spores are monolete and oval to circular in polar view and reniform

(concavo-convex) in lateral view. The laesura has distinctly raised lips for most of

its length and is about one-half the total length of the spore. The exine is thick and

robust and is uniformly and densely covered by regularly arranged, shallow

microfoveola. Shallow furrows may connect adjacent microfoveola. The overall

dimensions are very regular, approximately 80 x 60 w; length of laesura 40 yw,

exine 3 w, depth of foveola 0,5 uw, 11-16 foveola per 100 pw’.

Remarks

Harris (1965) does not make it clear how Microfoveolatosporis fromensis

(Cookson) Harris, 1965 differs from Microfoveolatosporis pseudodentatus

Krutzsch, 1959, the type species of the genus.

28 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

The genus Microfoveolatosporis has affinities with the genus Schizaea of the

Schizaeaceae. Spores of Schizaea pennula Swartz, an extant Columbian species,

are indistinguishable from M. fromensis (Murillo & Bless 1978: 356).

Distribution

Microfoveolatosporis is known from the Tertiary of both hemispheres. The

extant genus Schizaea is distributed predominantly in the Southern Hemisphere,

with two species occurring in southern Africa (Welman 1970). Microfoveolato-

sporis fromensis exits from the tropical African record at the Cretaceous—Tertiary

boundary (Salard-Cheboldaeff 1979).

Genus Cicatricososporites Pflug & Thomson,

in Thomson & Pflug, 1953

See discussion of Cicatricososporites and Schizaeoisporites in Jansonius &

Hills (1976: 468-469, 2530) and Srivastava (1971: 256).

Cicatricososporites sp.

Fig. 6D

Compare

Cicatricososporites norissii Srivastava, 1971: 257, pl. 1 (figs 5-8).

Description

The spore is alete or possibly monolete, with canaliculate to cicatricose

sculpturing. Odd ridges may bifurcate for a short distance and the sculpturing is

somewhat asymmetrical, resulting in a slight spiralling appearance. No laesura

was observed, but it may lie parallel to, and be almost indistinguishable from, the

grooves of the sculpturing. The length of the specimen illustrated is 60 pw.

Affinity

The genus Cicatricososporites has affinities with the genus Schizaea.

Srivastava (1971) stated that ‘Cicatricososporites norissii is comparable with

spores of the extant species Schizaea laevigata illustrated by Selling 1946’.

Distribution

Cicatricososporites norissii is common in the lower member of the Edmonton

Formation (Maastrichtian), Canada, where it is a prominent component of a flora

that Srivastava (1971) suggested grew under humid conditions. The distribution

of Cicatricososporites in the Palaeocene presents a strange disjunct pattern, which

is reflected in the present-day distribution of the genus Schizaea (see Table 2).

Only one specimen of Cicatricososporites sp. was observed at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 29

Alete spores

Genus Reticulatasporites Ibrahim, 1933

Ibrahim’s (1933) diagnosis of the genus Reticulatasporites was very general:

‘Spores without trilete mark, and with a measurable reticulate sculpture on the

spore wall; meshes up to [or as small as ? JJ] 1 «’ (Ibrahim 1933: 38, in Jansonius

& Hills 1976: 2362). It is clearly a rather insecure ‘portmanteau’ genus. Potonié &

Kremp (1954), quoted in Jansonius & Hills (1976), provided a more precise

diagnosis intended to accommodate a group of fungal spores.

The species described below conforms in its general features to Reticulata-

sporites (sensu lato), so that in the absence of a thorough revision of the genus and

its related forms it has been attributed to this genus, despite marked differences

from the type species of the genus.

Reticulatasporites grandis sp. nov.

Fig. 6E—-F

Etymology

The name of this species reflects the large size of the grain.

Description

The spores are atreme and spherical to subspherical—ellipsoidal. The whole

surface of the spore is covered by a large-meshed, quite regularly sized reticulum,

the muri formed apparently by steep folding of the outer skin. The muri

themselves are never straight but ‘wriggle’ across the surface. The lumina are

polygonal, mostly pentagonal, in shape. A pinnacle may be formed at the

junction of the muri. The diameter of the grains is 50-60 w.

Remarks

Reticulatasporites grandis may be more similar to other species of Reticulata-

sporites than it is to the type species of the genus. There is a superficial

resemblance between R. grandis and Retiperiporites piacabucuensis Herngreen

(1975b: 110, pl. 2 (fig. 5)), known from the Upper Senonian of Brazil.

Affinity

The affinities of Reticulatasporites grandis are open to speculation. It is

unlike lycopodiaceous spores. It is more likely that it has an affinity with a

bryophyte family, such as the Cleveaceae, whose members may also produce

atreme spores with large reticulate features (Boros & Jarai-Komlddi 1975).

Distribution

Reticulatasporites grandis is common in the Arnot samples. There is only

negative information about its wider distribution. It has apparently not been

recorded in Australian late Cretaceous and Tertiary sediments. A single specimen

30 ANNALS OF THE SOUTH AFRICAN MUSEUM

encountered in late Cretaceous sediments off the south-western Cape coast and

described as ‘Lycopodiumsporites facetus’ appears from the photomicrograph

(McLachlan & Pieterse 1978: 875, pl. 2 (fig. 1)) to be indistinguishable from the

present form.

Spores not assigned to genus

Forma A

Fig. 7C-D

Few specimens of this spore were seen and it was unclear whether the robust

verrucate structures occurred on both faces. It is probably verrucate on the distal

face alone, which would place it in the genus Distaverrusporites, known from the

late Cretaceous of Nigeria and Borneo (cf. Van Hoeken-Klinkenberg 1966: 43,

pl. 1 (fig. 6)). Equatorial diameter 50 w, height of verrucae 6 wp.

Forma B

Fig. 8G-—H

Few specimens of this spore were seen. Equatorial diameter 32-34 wp.

Reticulum only on distal face. Muri 2 wu high. The junction of muri is marked by a

truncated spinate process, which is an additional 3—4 w higher than the muri. This

spore can probably be assigned to the genus Retitriletes, which has lycopodiaceous

affinities.

Forma C

Fig. 7E-F

Equatorial diameter 50 w. The spore is trilete. Laesurae are straight and two-

thirds the radius of the spore. Distal face is hamulate. Proximal face is covered by

a perine, loosely attached to the exine and tightly folded to form a fine rugulate—

hamulate pattern. Exine 3 w. This spore could perhaps be assigned to Hamulati-

sporis Krutzsch, 1959 (a subgenus of Camarazonosporites), which is known from

late Cretaceous and Eocene deposits in Europe (Azema & Ters 1971: 272).

Forma D

Fig. 7A-B

This is a very large trilete microspore with a massive perinous layer, up to

20 mw thick on the distal face and the equatorial region. This layer is coarsely and

jaggedly verrucate or fossulate. The proximal face is subpsilate. The laesurae are

thin and straight, or with a slight curve, and almost reach the equator. They are

placed on the apex of raised triangular ridges running the length of the radial

areas. This form distinctly resembles the spores of the extant dwarf tree-fern

genus Lophosoria (family Lophosoriaceae) as illustrated in Murillo & Bless

(1974: 252-253). This neotropical family is sometimes included in the

Cyatheaceae.

PALYNOLOGY OF THE ARNOT PIPE

eg Ae or eee

404

Fig. 7. A-B. Forma D. C-D. Forma A. E-F. Forma C (Hamulatisporis).

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 8. A. Araucariacites australis. B. Araucariacites sp. C-F. Zonalapollenites sp. B

(Cingulatipollenites). G—H. Forma B.

PALYNOLOGY OF THE ARNOT PIPE 35

Pollen of Coniferae

Inaperturate pollen

Genus Araucariacites Cookson ex Couper, 1953

Araucariacites australis Cookson ex Couper, 1953

Fig. 8A

Granulonapites (Araucariacites) australis Cookson, 1947: 130, pl. 13 (figs 1-4).

Araucariacites australis Cookson, 1947. Balme, 1957: 31, pl. 7 (figs 81-82). Kemp & Harris, 1977:

25, pl. 4 (figs 15-16).

Description

The pollen grains are large, always flattened and crumpled; inaperturate and

with a thin, finely granulate exine. Diameter 50-60 pw, exine 0,5 pw.

Affinity

See discussion in Cookson (1947: 130) and Kemp & Harris (1977: 25). The

affinity of A. australis is with the Araucariaceae and probably the genus

Araucaria.

Distribution

Araucariacites australis is common at Arnot, reaching relative abundances of

3 per cent. This species has a world-wide distribution in the Mesozoic, but is

mainly restricted to the Southern Hemisphere in the Tertiary. So far as is known

it is not present in the Neogene of the African sub-continent (Thiergart er al.

1963; Coetzee 1981). It is common in lower Cretaceous sediments off the east

coast of southern Africa (Scott 1976; McLachlan & Pieterse 1978), but less

common in the interior (Scholtz & Deacon 1982) and south-western Cape in

Cretaceous sediments. Together with other inaperturate forms, it dominates early

Palaeocene or late Cretaceous assemblages from the Gamoep area (unpublished

data).

Araucariacites sp.

Fig. 8B

Compare

Inaperturopollenites limbatus Balme, 1957: 31, pl. 7 (figs 83-84).

Balmeiopsis limbatus (Balme) Archangelsky, 1977: 122-126, pl. 1.

Araucariapollenites laffittei Reyre, 1973: 157, pl. 35 (figs 3-4).

Description

Outline oval or subcircular, exine 2-3 yw thick and granulate, diameter

57-74 w.

Discussion

The exine is much thicker than that of Avraucariactes australis but the

granulate sculpturing is very similar. Few specimens were seen in the samples

from Arnot, and although preservation of all specimens was excellent, doubt

34 ANNALS OF THE SOUTH AFRICAN MUSEUM

about its morphology remains. Possibly what is here described as a thick exine, is

rather an equatorial crassitude or a particular concentric folding pattern of larger

specimens of the thinner-exined A. australis. Inaperturopollenites limbatus

Balme, 1957, is large, robust, granulate and inaperturate. Archangelsky (1977)

has recently instituted the genus Balmeiopsis for larger spherical, granulate grains

with an equatorial crassitude and an irregular aperture or thinning of the exine at

one pole. Reyre (1973) erected Araucariapollenites on the rather tenuous basis of

SEM-observed ultrasculptural exine features. Since few specimens were observed

at Banke and an irregular aperture was not noted, a definitive description of the

grain is left for later research. Palynomorph assemblages from other crater-lake

deposits in the same area, but somewhat earlier in time, consistently contain

A. australis and a larger and more robust morphotype.

Affinity

Araucariacites sp. is probably an extinct species of Araucaria. Balmeiopsis is

found in association with twigs and cones associated with Brachyphyllum-type

foliage, a leaf genus with affinity to the Araucariaceae (Archangelsky 1977).

Reyre (1973) compares Araucariapollenites to the pollen of Araucaria araucana

(cf. A. araucana Heusser, 1971: 12, pl. 8 (fig. 55)).

Distribution

Araucariacites sp. occurs in the Australian lower Cretaceous where it is

always rare (Balme 1957: 31; Burger 1973: 100). Balmeiopsis is recorded from

lower Cretaceous sediments of South America and Canada (Archangelsky 1977)

and Araucariapollenites from Mesozoic sediments of north Africa (Reyre 1973).

Monosaccate pollen

Genus Zonalapollenites Pflug, in Thomson & Pflug, 1953

See discussion in Dettmann (1963: 99, under Tsugaepollenites) and in

Jansonius & Hill (1976: 3093, 3265).

Zonalapollenites sp. A

Fig 9A—D

Description

The corpus is biconvex or boat-shaped in transverse section and the grain is

perisaccate in the equatorial—subequatorial region. The amb is roundly rectangu-

lar to broadly elliptical. Distal and proximal faces appear to have fused so as to

create a thick, laminated main body, which is granulate and rugose to

indeterminately sculptured. Initial folds in the exoexine on both the proximal face

(the face on to which the saccus overlaps) and the distal face mark the attachment

of the saccus. The saccus is formed from a relatively thick exoexine, which is

sometimes deeply folded in a radial direction and is distinctly less well developed

at the opposing poles of the longest axis of the amb. It would appear that the zone

of attachment of the saccus to the main body is subequatorial on the proximal and

PALYNOLOGY OF THE ARNOT PIPE 35

Fig. 9. A-D. Zonalapollenites sp. A. E. Lygistepollenites sp. F-1. Podocarpidites sp.

36 ANNALS OF THE SOUTH AFRICAN MUSEUM

perhaps also the distal face, resulting in a characteristic subequatorial dark ring

when the grain is viewed from the polar position. This effect is caused by the

density of the sporopollenin in the equatorial to subequatorial zone. Total

diameter 33-49 w, saccus width 9-17 w, diameter of the main body 28-38 yp,

thickness of main body 3—4 pw, height of the whole grain 22 wp.

Discussion

This species is similar to those described as Zonalapollenites segmentatus

(Balme 1957: 33, pl. 9 (figs 93-94)) and Tsugaepollenites segmentatus (Dettmann

1963: 101, pl. 24 (figs 6f, 11-16)) but differs clearly from the description of the

latter in that the present species does not have polar vesiculae. The present

species also differs in the characteristic wide, dark subequatorial (taking the grain

as a whole) ring. It is very difficult to photograph in transverse section. It must be

pointed out that the original generic diagnosis specifies that the “saccus’ is in fact a

velum formed by a fibrous baculate extension of the exine. Dettman’s (1963)

diagnosis of the genus mentions only an equatorial saccus and both Zonala-

pollenites sp. A and Zonalapollenites sp. B, described below, fit this more recent

diagnosis.

Affinity

Dettmann (1963: 100) quotes other authors in support of a coniferous

affinity — possibly to the genus Tsuga. Muller (1968) quotes Gamerro (1965) who

suggests a podocarpaceous affinity for Zonalapollenites.

Distribution

The genus is present world-wide in sediments of Jurassic to Palaeogene age.

It is rare in sediments of Cretaceous age off the southern African coast.

Zonalapollenites sp. A is rare at Arnot.

Zonalapollenites sp. B

Compare

Cingulatipollenites aegyptiaca Saad & Ghazaly, 1976: 449, pl. 13 (figs 3-6).

Description

The structure is complex. The grain is biconvex to flat. The amb is

subcircular to broadly elliptical. Monosaccate equatorially with the limbatus

marked by a distinct, sharp, irregular line. The saccus is of medium width and it is

not prominently folded in a radial or any other direction, but is rather

characterized by superficial pliae. The saccus is robust and may terminate

equatorially in a crassitude. The exine of the main body is relatively thin and

granulate proximally and distally. The outline of the grain is irregular and the

saccus may overfold the proximal surface, resulting in a rough frill-like line

internal to the limbatus. On occasional grains a trilete fold may mark laesurae.

The large size range makes it likely that more than one species is present.

Diameter 36-70 yw, width of saccus 6-12 wp.

PALYNOLOGY OF THE ARNOT PIPE a7

Discussion

The grains are different from those of Zonalapollenites sp. A in having a

relatively much smaller saccus, which is not folded in a radial direction. The

saccus in the present species also does not so regularly or prominently overfold

the proximal surface. The exine in the present species is granulate, while in

Zonalapollenites sp. A it is granulate—rugose. Although the outline of the grain is

irregular it does not show the undulate outline of Zonalapollenites dampieri, nor

has the overfolding of the saccus of the proximal face been mentioned as a

characteristic of Z. dampieri (Balme 1957: 32, pl. 8 (figs 88-90); Dettmann 1963:

100, pl. 24 (figs 1-5)). In other respects the present species is similar to

Z. dampieri.

Saad & Ghazaly (1976) have described Cingulatipollenites aegyptiaca from the

Nubia Sandstones of North Africa, which is almost certainly conspecific with

certain of the forms encountered in this study. However, some of the present

specimens do appear clearly saccate, rather than cingulate, so that the genus

Zonalapollenites is preferred in the present context. As indicated the size range and

morphological variability make it likely that more than one species is present, and

perhaps cingulate as well as saccate forms. Further study of these forms is required.

Affinity

Zonalapollenites sp. B is presumably coniferous, but there is no further

information on its specific affinities.

Distribution

Zonalapollenites sp. B is common at Banke. The genus is known world-wide

from the Jurassic to the Palaeogene. Cingulatipollenites is present in Jurassic to

Upper Cretaceous assemblages from the Nubia Sandstones, North Africa (Saad

& Ghazaly 1976).

Disaccate pollen

Genus Lygistepollenites Stover & Evans, 1973

Lygistepollenites sp.

rice IE

Compare

Dacrydium cupressinum Couper, 1953, pl. 4 (fig. 35).

Description

The grain is disaccate; the corpus is circular and covered proximally and

laterally with a wide layer of steep folds forming a rugulate pattern. The sacci are

small and half pendent on the distal face. The sacci seem to be formed merely by

larger folds of the outer skin. They are folded in a radial direction and there is no

distinct zone of attachment. The distal face bears less robust sculptural elements

and the sulcus, wide in the centre and narrowing towards the equatorial margins,

is clear.

38 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Only one specimen in a good state of preservation was observed. The grain

lacks prominent proximal protuberances between the body and the proximal

roots of the sacci and thus does not belong to the genus Phyllocladidites Cookson

ex Couper, 1953, despite its small sacci. The grain is similar to grains of the extant

species Dacrydium cupressinum (Pocknall 1981: 70, figs 2a—e), although its sacci

may not be as well developed.

Affinity

The affinity of the genus Lygistepollenites is with the genus Dacrydium

(Section B) of the Podocarpaceae.

Distribution

Only one grain of Lygistepollenites sp. was observed at Arnot. The genus

Lygistipollenites is known from the Oligocene to the present in New Zealand. It

has not been recorded in late Cretaceous sediments from the interior of the

African subcontinent (Scholtz & Deacon 1982) nor off the Cape (McLachlan &

Pieterse 1978), but it is probably present in the Neogene (?) in the southern

(Thiergart et al. 1963) and south-western Cape (J. A. Coetzee, Institute for

Environmental Sciences, University of the Orange Free State, pers. comm.). It is

not recorded from late Neogene sediments from Burundi (Sah 1967).

Genus Podocarpidites Cookson ex Couper, 1953

Podocarpidites sp.

Fig. 9F-I

Compare

Disaccites grandis Cookson, 1953: 47, pl. 2 (fig. 41).

Pityosporites grandis (Cookson) Balme, 1957: 36, pl. 10 (figs 110-111).

Alisporites grandis (Cookson) Dettman, 1963: 102, pl. 25 (figs 1-5). Haskell,

1968: 217, pl. 1 (figs 1-2).

Description

The corpus is circular in polar view and the exine is thick. The outline of the

corpus is sometimes difficult to see. The sacci are large, semicircular and slightly

wider than the corpus. On the distal face their roots are clearly marked, spaced

wide apart and parallel, and outline a correspondingly broad tenuitas. The

reticulum of the sacci is coarse and often discontinuous towards the margins of the

sacci. The grain is sometimes apparently collapsed, in which case the tenuitas

appears narrower and tending towards fusiform while the shape of the grain

becomes oval. The preservation is usually poor. Length of the expanded grain

88-119 w, diameter of the corpus 60-63 pw, length of the sacci 68—75 ww, breadth

of the sacci 25-32 yw, distance between the zones of attachment on the distal face

25-33 p, exine of irregular height, 4-8 wp.

PALYNOLOGY OF THE ARNOT PIPE 39

Discussion

Haskell (1968: 217) provides a satisfactory description on the range of

variation in Alisporites grandis. The dimension of the grains that he measured are

very similar to those of Podocarpidites sp. He described two states for the

form—a diploxylonoid and a haploxylonoid state—which may correspond to the

‘expanded’ and ‘collapsed’ states described here. Despite regarding the present

form as conspecific with A. grandis (Cookson) Dettmann, 1963, as described by

Haskell (1968), and since the ‘expanded’ or diploxylonoid state (which shows

non-Alisporites-like characteristics) is most common at Banke, it was not

considered justified to place it in the genus Alisporites. It is beyond the scope of

the present paper to propose new combinations, and it is therefore merely noted

that the genus Alisporites Daugherty, 1941, has been redefined (Jansonius 1971;

Jansonius & Hill 1976: 68-69) and, in the opinion of the present author, the

diploxylonoid state of Podocarpidites sp. and of A. grandis as redescribed by

Haskell (1968) does not permit the inclusion of these species in the revised

diagnosis of the genus Alisporites. Jansonius (1971) also suggested a pterido-

spermous affinity for Alisporites, while the morphology of the present form

suggests a podocarpaceous affinity. The present form is probably comparable to

that described as Podocarpidites sp. by Sah (1967: 44, text-fig. 13, pl. 4 (fig. 11))

from the Neogene of Burundi.

Affinity

The affinity of Podocarpidites sp. is probably with the Podocarpaceae,

section Eupodocarpus (which includes the African species Podocarpus latifolius,

P. elongatus and P. henkelii) or section Stachycarpus (A. R. H. Martin 1959;

Pocknall 1981). On overall size alone the New Zealand members of what has

been regarded as the most primitive section of the family, Stachycarpus (Bucholtz

& Gray, 1948), compare most closely with the present form. The rarity and state

of preservation of the specimens precludes finer morphological comparisons.

Distribution

Alisporites grandis is known from the upper Jurassic and lower Cretaceous

strata in Australia and Canada (Haskell 1968) and from Cretaceous strata off

South America (Archangelsky & Gamerro 1967). It is common in the Australian

lower Cretaceous. It is not recorded in Cretaceous sediments of DSDP 361 off the

south-western Cape coast (McLachlan & Pieterse 1978), nor is it present in the

Knysna lignites (Thiergart et al. 1963). It is present in the Neogene of central

Africa (Sah 1967), where it is very rare. Podocarpidites sp. is rare at Arnot.

Podocarpidites kamiesbergensis sp. nov.

Fig. 1OA—D

Etymology

This species is named after the nearby Kamiesberg Mountains.

40 ANNALS OF THE SOUTH AFRICAN MUSEUM

eal

”

Fig. 10. A-D. Podocarpidites kamiesbergensis sp. nov. E-J. Podocarpidites riembreekensis

sp. nov. K-L. Arecipites sp. B. M-—P. Arecipites sp. A.

PALYNOLOGY OF THE ARNOT PIPE 4]

Description

Small, disaccate pollen grains, the corpus trapeziform in shape, but

somewhat arched proximally. The semi-hemispheric sacci are attached laterally at

a low angle. The grain is often not fully expanded, in which case the sacci appear

more pendent. The exine is relatively thick proximally and laterally, and

vermiculate to foveolate. The distal tenuitas is relatively broad and parallel-sided

and the infrareticulation of the sacci is robust, clear and mostly perfect. In polar

view the sacci are as broad as the corpus. Corpus circular in polar view. Outline of

the sacci regular. Total length of expanded grain 36—41 uw; height of corpus in

lateral view 20 w, width of corpus 23-26 uw, length the same; breadth of sacci

10 pw, depth 16 pw, length of sacci 20-23 yw, distance between lines of attachment

of sacci on the distal face 8-10 yp.

Discussion

Podocarpidites kamiesbergensis is smaller in size than the smaller species of

Podocarpidites described in the available literature, such as P. congoensis Sah

(1967: 43, pl. 4 (figs 5-6, 9-19)). Podocarpidites kamiesbergensis may be the same

as P. knysnanus Thiergart, Frantz & Raukopf, 1963, described from the Knysna

lignites, but this form is inadequately characterized and illustrated (Thiergart er

al. 1963).

Affinity

The affinities of P. kamiesbergensis are with the Podocarpaceae, especially

the section Afrocarpus (including Podocarpus gracilior and P. falcatus) as defined

by A. R. H. Martin (1959). It would be difficult to distinguish between the pollen

of P. falcatus and Podocarpidites kamiesbergensis.

Distribution

Podocarpidites kamiesbergensis is common at Arnot, reaching a relative

abundance of 5 per cent. It is not present in the late Cretaceous sediments from

the interior (Scholtz & Deacon 1982) or off the south-western Cape (McLachlan

& Pieterse 1978), but is possibly present in the Neogene (?) Knysna lignites.

However, the description of P. knysnanus Thiergart, Frantz & Raukopf, 1963,

does not permit identification with the present species.

Podocarpidites riembreekensis sp. nov.

Fig. 10E—J

Etymology

This species is named after the nearby farm Riembreek, which is also the site

of several crater-lake deposits.

Description

Medium-sized disaccate pollen grains. The corpus has a rounded rhomboidal

shape in polar view and is trapeziform in side view. The sacci are laterally

attached at a low angle and are rigid, hemispherical and slightly wider than the

42 ANNALS OF THE SOUTH AFRICAN MUSEUM

corpus in polar view. The infrareticulum of the sacci is greatly reduced so that

only isolated sections of muri remain. The proximal and lateral surfaces of the

corpus are covered by sharply defined, densely packed rugulate to verrucate or

vermiculate structures, and the exine thickens in the proximal area to form a

distinct cappa. Proximally each saccus is attached to the corpus by two

crassitudes, which are not always prominently protuberant. Such protuberances

are not so well developed around the remaining lateral and distal zones of

attachment, but are developed enough to form a thick rugose collar that

constitutes the roots of the sacci. The roots are marked, but least robust on the

distal face where the sacci are separated by a wide, parallel-sided tenuitas.

Total length of grain 40-64 w; width of corpus 23-33 mw, length of corpus

25-35 mw; length of sacci 25-39 w, depth of sacci approximately 14 w, breadth of

sacci approximately 25 w, exine of sacci approximately 1 wp.

_ Discussion

The grain has very distinctive morphology. The infrareticulum of the sacci is

so reduced that it may be entirely absent over large areas. The exine of the sacci is

relatively thick. In polar view the four proximal crassitudes are marked and

produce the characteristic rhomboidal shape of the corpus. The proximal and

lateral sculpturing is distinct and robust, and is truncated as the distal surface is

approached. The proximal protuberances are part of the robust collar that

attaches the sacci to the corpus, and differ from the more localized structures

described for Phyllocladidites. The much larger sacci also distinguish this species

from Phyllocladidites.

Affinity

The affinity of Podocarpidites riembreekensis is with the Podocarpaceae, but

perhaps not with any extant genus.

Distribution

Podocarpidites riembreekensis is common at Arnot. As far as is known this

form has not been recorded elsewhere.

Pollen of Angiospermae

Monoaperturate pollen

Monocolpate pollen

Genus Arecipites Wodehouse, 1933

‘The genus Arecipites Wodehouse, 1933 was emended [sic] by Anderson

(1960) to include only those reticulate monosulcate pollen grains which have,

among other characters, lumina less than 0,5 wu in diameter. Reticulate

monosulcate forms whose lumina width exceeds 0,5 w were referred by Anderson

to the genus Liliacidites Couper, 1953’—Chmura (1973: 104). This procedure has

been generally accepted and is the one followed here.

PALYNOLOGY OF THE ARNOT PIPE 43

Arecipites plectilimuratus Chmura, 1973

Fig. 11A—B

Arecipites plectilimuratus Chmura, 1973: 104, pl. 21 (figs 1-3).

Description

Monosulcate; elongate—ellipsoidal in polar view. The sulcus extends the

whole length of the grain and may even transgress the ends of the grain. The

margins of the sulcus are faint and irregular since the exine thins as the sulcus is

approached. The sulcus is open and, despite the irregularity, more or less

parallel-sided for its whole length. The exine is relatively thick and the reticulum

is distinct and uniform over the whole of the grain, except when the sulcus is

approached and the reticulum becomes indistinct. Muri are approximately the

same width as the lumina and mostly duplibaculate. Length of illustrated grain

40 pw, width 28 pw, exine 2 yp.

Discussion

For comparison with other species of Arecipites see Chmura (1973: 104). The

genus is a generalized morphological type and no specific identity of the plants

involved need be supposed.

Affinity

The affinity of Avicipites plectilimuratus is with a broad monocotyledonous

group including the Amaryllidaceae, Iridaceae and Liliaceae. The pollen of

southern African Monocotyledonae is not sufficiently well known to enable one

to suggest closer affinities with any taxa of the local flora.

Distribution

Arecipites plectilimuratus is rare at Arnot. It is also rare in the late

Cretaceous of California.

Arecipites sp. A

Fig. 1OM-—P

Compare

Arecipites reticulatus (van der Hammen) Anderson, 1960: 18.

Description

Small, monosulcate pollen, elongate—oval with rounded ends. The sulcus is

long, has no margo and reaches the ends of the grain. The exine is relatively thick

and clearly differentiated into a nexine and a tectate sexine. The surface of the

grain is uniformly covered by microscrobiculi. Length of the grain 23-25 yw, exine

1—2 p thick, scrobiculi 0,2 pw.

Fig. 11.

ANNALS OF THE SOUTH AFRICAN MUSEUM

A-B. Arecipites plectilimuratus. C-D. Liliacidites sp. B. E-F. Clavatipollen-

ites spp. (SEM). G-L. Clavatipollenites sp. A. M-N. Clavatipollenites sp. B.

O-Q. Clavatipollenites sp. C. R-T. Monocolpopollenites sp. B.

PALYNOLOGY OF THE ARNOT PIPE 45

Discussion

Arecipites sp. A is a small, robust, distinctive grain that is dissimilar to

A. plectilimuratus. It differs from A. reticulatus in its finer scrobiculi and thicker

exine.

Affinity

Arecipites sp. A has monocotyledonous affinities.

Distribution

Arecipites sp. A is rare at Arnot.

Arecipites sp. B

Fig. 1OK-L

Description

The grain is ellipsoidal with a thin and indistinct colpus, which runs the

length of the grain. The exine is relatively thick and the sexine and nexine layers

are clearly differentiated. The sexine is psilate to indeterminately sculptured and

appears finely columellate. Dimensions of illustrated specimen 38 X 26 w, exine

1-2 p.

Affinity

No information exists regarding the affinity of Arecipites sp. B.

Distribution

Arecipites sp. B is rare at Arnot.

Genus Liliacidites Couper, 1953

Liliacidites sp. A

Fig. 12E—H

Description

The grains are ellipsoidal to elongate—ellipsoidal in polar view. The sulcus in

expanded grains is wide with parallel sides, and stretches the length of the grain.

The meshes of the complex reticulum become smaller as the sulcus is approached

until a margo is formed by a narrow zone of unbroken exine. The exine may also

become thinner towards the colpus, but this was not definitely established.

Elsewhere the reticulum forms a complex, irregular, non-perfect pattern and is

simplicolumellate. The exine is relatively thick and two layers are clearly

distinguishable. Sexine and nexine are of similar height. Length of grain 29-36 p,

width of expanded grain in polar view 30 uw, sulcus width 3,5-5 w, exine

approximately 2 w, lumina 1-3 wp.

46 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

Liliacidites sp. A appears to be similar to those described as Liliacidites

intermedius Couper, 1953, but is distinguished from that species by the

characteristic reduction of the reticulum in the vicinity of the colpus, and in that

the reticulum of the present species is not reduced towards the ends of the grain.

Affinity

The affinity of Liliacidites sp. A is perhaps with the genus Chamaedorea of

the Palmae. Similar forms are also found in the Butomaceae and Liliaceae. The

extant genus Chamaedorea contains rather unusual small, reed-like palms found

in tropical deciduous thickets (Lozano-Garcia 1979).

Distribution

Liliacidites sp. A is common at Arnot.

Liliacidites sp. B

Fig. 11C-—D

Description

Monocolpate, irregular elongate—ellipsoidal. Narrow sulcus with distinct

margo stretches the full length of the grain. The tectum is imperfect.

Crassisexinous. The reticulum is irregular, imperfect and robust, so that the

sculpturing appears reticulate-rugulate. The columellae may be horizontally

elongated so that the sculpturing gives the appearance of broken stretches of

muri. Length of grain illustrated 36 w, width 22 pw, exine 3 w.

Discussion

Liliacidites sp. B is most unusual in the reduction in height of the columella

and a corresponding increase in the depth of the tectum.

Affinity

The affinity of Liliacidites sp. B is perhaps with the Liliaceae (cf. Lilium

longiflorum (Huang 1972: 266, pl. 173)).

Distribution

Liliacidites sp. B is very rare at Arnot.

Genus Clavatipollenites Couper, 1958

See discussions in Dettmann (1973: 11) and Kemp & Harris (1977: 55). The

genus is well represented at Banke with at least three forms being present, one of

which is common.

PALYNOLOGY OF THE ARNOT PIPE 47

Clavatipollenites sp. A

Fig. 11G—L

Compare

Clavatipollenites sp. Dettmann, 1973: 11, pl. 2 (figs 8-11).

Description

The grains are monocolpate and subspheroidal to slightly ellipsoidal—

spherical. The colpus is usually ulcerate with a broken and irregular margin in the

sexine. It is irregular in outline and usually more or less isodiametric. Occasional

grains have a clean, straight margin in the sexine (Fig. 111) and others appear

trichotomosulcate (Fig. 11J). Sexine and nexine are about the same height. The

reticulum is perfect and the muri simplicolumellate. The swollen heads of the

columellae and thus ‘lumpiness’ of the muri can be seen in the SEM

photomicrographs (Fig. 11E—F). Further SEM work may confirm whether the

‘lumpiness’ (Kemp & Harris 1977) or ‘beaded’ characteristics (Coetzee 1981) of

the exine can be of taxonomic significance. It would appear from the limited SEM

work done in the present study that in Clavatipollenites sp. A the ‘beadedness’ is

caused by suprategillar microconi mounted on the branches of the reticulum,

while in Clavatipollenites sp. C (described below) the ‘lumpiness’ of the muri may

be caused by the swollen heads of the columellae alone. Length of grain usually

between 22-27 uw, but occasional grains up to 31 mu, exine 1,5-3 w, lumina

approximately 1 mw, sulcus usually in the order of 5—8 w diameter, but may be

longer.

Affinity

See discussions in Dettmann (1973: 11), Kemp & Harris (1977: 56) and

Muller (1981: 9). Affinity of the genus Clavatipollenites is probably with the

Chloranthaceae (Doyle 1969).

Distribution

The distribution in time and space and a suggested pattern of extinction of

the Clavatipollenites—Ascarina complex is discussed in Muller (1981: 9-12). Their

suggested pattern of extinction needs to be revised since Clavatipollenites has

been recorded at Arnot and in the Neogene in the south-western Cape (Coetzee

1981). Observations from the Botswana region (Scholtz & Deacon 1982),

however, confirm that in late Cretaceous assemblages dominated by Ephedripites

(and Cretacaeisporites), Clavatipollenites is not present. Clavatipollenites sp. A is

common at Arnot.

Clavatipollenites sp. B

Fig. 1LM-N

Description

Monocolpate, subspherical to ellipsoidal—spherical. Sulcus relatively smaller

than in Clavatipollenites sp. A and more circumscribed. Exine thick and

48 ANNALS OF THE SOUTH AFRICAN MUSEUM

crassinexinous. Recticulum relatively fine. Length 24 uw, width 19 w, nexine

approximately 2 w, sexine 1 w, lumina 1 wp.

Discussion

Clavatipollenites sp. B is distinguished from Clavatipollenites sp. A by the

prominent dense nexine, relatively thin sexine, and fine reticulation.

Distribution

Clavatipollenites sp. B is rare at Arnot.

Clavatipollenites sp. C

Fig. 110-Q

~ Description

Monocolpate, ellipsoidal—spherical with a long closed colpus, which, as usual

for Clavatipollenites, has ragged broken edges in the sexine. The exine is

relatively thick with sexine and nexine about the same height. Markedly robust

columellae support a perfect reticulum. Columellae are only placed beneath wide

areas of the tectum formed at the junction of individual branches of the tectum.

Length of grain 32 uw, width 26 w, exine 3-4 yw, lumina 1-2 p.

Discussion

Clavatipollenites sp. C is clearly distinguishable from the former two species

by its greater size, long sulcus, and robust columellae and tectum.

Distribution

Clavatipollenites sp. C is rare at Arnot.

Genus Monocolpopollenites Pflug & Thomson,

in Thomson & Pflug, 1953

See discussion in Nichols et al. (1973) and Jansonius & Hill (1976: 1691).

Monocolpopollenites sp. A

Fig. 12A—D

Compare

Monocolpopollenites sp. Jardiné & Magloire, 1963: 211-212, pl. 8 (figs 31-32).

Description

Monocolpate, subcircular to ellipsoidal in polar view and ellipsoidal in

equatorial view. The colpus has a complex structure; it is parallel-sided and

reaches the ends of the grain. The lips of the colpus are folded inwards and

PALYNOLOGY OF THE ARNOT PIPE 49

Fig. 12. A-D. Monocolpopollenites sp. A. E-H. Liliacidites sp. A. I-J. Forma E

(monocolpate pollen). K-—M. Milfordia hypolaenoides. N. Milfordia sp.

50 ANNALS OF THE SOUTH AFRICAN MUSEUM

thickened, and on the inner margins of the lips a scabrate row of sculpturing is

formed, consisting perhaps of closely packed, small verrucae. This is a very

characteristic feature. The exine is thick, tectate and psilate, and a finely punctate

perinous membrane envelops the grain. This layer sometimes closely adheres to

the surface of the grain and at other times separates from the exine. Few

specimens were observed, but from the size range it is possible that two species

may be present. Length 26-39 w, width 18-34 uw, exine 2,5—3 w, width of colpus

measured from the outer margins of the interior sculptured lips 5-8 w.

Affinity

The affinity of Monocolpopollenites sp. A is not known.

Distribution

Monocolpopollenites sp. A is rare at Arnot. Monocolpopollenites sp. Jardiné

& Magloire, 1963, is known from Turonian to Maastrichtian time ranges from

Senegal and the Ivory Coast.

Monocolpopollenites sp. B

Fig. 11R-T

Description

Grains monocolpate, rarely trichotomosulcate; ellipsoidal to subcircular.

The colpus stretches the length of the grain, is parallel-sided and relatively wide.

The lips are raised and decorated with short folds or verrucae. The ends of the

colpus are abruptly truncated and edged with verrucae or bits of exine. The exine

is relatively thin and is sparsely and irregularly decorated with isolated spinosa.

Length of grain 30-34 uw, width 20-23 uw, width of colpus approximately 4 uw,

exine approximately 1 wp.

Discussion

Monocolpopollenites sp. B is distinct from Monocolpopollenites sp. A in

having a much thinner exine and lacking a perinous layer. The latter may be a

weak criterion considering that only a few grains were observed. The sparse, but

always present, isolated spinosa of Monocolpopollenites sp. B are also distinctive.

Affinity

The affinity of Monocolpopollenites sp. B is perhaps with the Palmae,

although as far as could be ascertained no extant Palmae display similar

sculpturing or end-aperture morphology (Sowumni 1972; Kedves 1980).

Distribution

Monocolpopollenites sp. B is rare at Arnot.

PALYNOLOGY OF THE ARNOT PIPE Sl

Monoporate pollen

Genus Milfordia Erdtman, 1960

Milfordia hypolaenoides Erdtman, 1960

Fig. 12K—M

Milfordia hypolaenoides Erdtman, 1960: 46, pl. 1 (fig. a). Martin, 1973: 37, figs 163-165.

Description

The pollen is monoaperturate and spherical to subspherical. The pore is

relatively large, circular or elliptical, and ulcerate, with the margins ragged and

often with loose pieces of exine in the mouth. The pollen has a slightly irregular,

undulating outline and the exine is relatively thick, scrobiculate and possibly

finely fossulate. Diameter of grain 25-32 « on SEM photomicrographs, 31-43 yu

on the light microscope; aperture diameter 11-20 w, occasionally smaller and

clotted with bits of exine; exine 1-2 wp.

Discussion

Although Elsik (1968: 313) and Partridge (in Stover & Partridge 1973: 262)

proposed generic diagnoses for types of restionaceous pollens that would include

both the smaller, porate or graminoid-type aperture and the larger centro-

lepidoid-type aperture (Chanda 1966), there is little value in using such a broad

concept on a specific level, as Partridge has done in Milfordia homeopunctata

Partridge, in Stover & Partridge, 1973. Since Chanda’s work (1966) it has been

clear that a distinction between the two types is significant in terms of the

phylogeny and plant geography (see Johnson & Briggs 1981: 458).

The normal aperture in the Arnot specimens of Milfordia hypolaenoides is

relatively large and ulcerate and clearly of the centrolepidoid type (Chanda 1966).

The aperture is similar to the ‘Hypolaena’-type illustrated in Couper (1960: 62,

pl. 9 (figs 26—27)) and H. A. Martin (1978: 191, pl. 7 (fig. X)) although,

unfortunately, as Muller (1981: 105) points out, only one species of the genus

Hypolaena has such a pore.

Erdtman (1938) and Chanda (1966) have suggested, on the basis of the

pollen morphology of extant species, an evolutionary sequence beginning with

the ‘primitive’ Centrolepidaceae, through those Restionaceae with centrolepidoid

apertures and the Restionaceae with graminoid apertures, to the ‘advanced’

Flagellariaceae and Poaceae (Chanda 1966: 396). Following the work of Hochuli

(1979), Muller (1981: 105) has noted the importance of the ‘relatively small

rounded—porate aperture with a more or less irregular margin and an indistinct

annulus which is transitional between the centrolepidoid and graminoid aperture

type’ in the early fossil record of the Restionaceae. This type of aperture has been

named the Restio subverticillatus-type by Muller (1981) and corresponds to the

‘Restio’-type of H. A. Martin (1978: 191, pl. 7 (fig. W)). The latter is once again a

Sy ANNALS OF THE SOUTH AFRICAN MUSEUM

rather unfortunate choice of name by Martin as the R. subverticillatus-type does

not include many members of the genus Restio (Chanda 1966).

It is clear that the evolutionary sequence proposed by Erdtman (1938) and

Chanda (1966) is not supported by the fossil record. All three apertural states of

the Restionaceae (graminoid, centrolepidoid, and transitional) are at present

known from late Cretaceous or Palaeocene sediments (Hochuli 1979; Salard-

Cheboldaeff 1979; Muller 1981; present study). It seems, however, that none of

the early forms show the extreme development of either centrolepidoid or

graminoid apertures that can be found in some extant species. In particular the

sharply protruding graminoid apertural state is as yet known only from possibly

early Neogene sediments (Thiergart et al. 1963).

The present evidence, confirming the early existence of the centrolepidoid

apertural state, indicates that the phylogenetic relationships inferred by Johnson

. & Briggs (1981) for the Restionaceae may need revision.

Affinity

Chanda (1966) and Ladd (1977) discuss the morphology of the pollen of the

Centrolepidaceae, Restionaceae and Flagellariaceae. Most modern Australian

species of Restionaceae have the centrolepidoid (Chanda 1966) or ‘Hypolaena’-

type apertures (Couper 1960; H. A. Martin 1978), while almost all southern

African species have the graminoid type (Chanda 1966). The present form,

Milfordia hypolaenoides, is therefore morphologically similar to extant Australian

Restionaceae rather than to the majority of southern African species. However,

the pollen of one southern African genus, Thamnochortus (Chanda 1966;

H. P. Linder, Bolus Herbarium, University of Cape Town, pers. comm.), is

comparable to the present form. Thamnochortus is unusual in a number of other

respects amongst the southern African Restionaceae, and the indications from

taxonomic studies are that it has had a long, isolated evolutionary history in the

subcontinent (H. P. Linder, pers. comm.). Milfordia hypolaenoides is also similar

to the pollen of Centrolepsis and Gaimardia of the Centrolepidaceae, but because

of the modern distribution of this family, an affinity between it and

M. hypolaenoides is considered unlikely.

Distribution

Milfordia hypolaenoides Martin, 1973, has been recorded from the late

Cretaceous of North America (Jarzen 1978) and in the Lower Palaeocene of

Europe where it continues into Miocene time ranges (Muller 1981). In Australia

it may be present from the lower Eocene (Stover & Partridge 1973) and is

common in younger sediments. Restionaceous forms were not recorded in late

Cretaceous sediments off the south-western Cape coast (McLachlan & Pieterse

1978). However, typically southern African Restio-type graminoid forms only

were recorded, and in great abundance, from the Neogene (?) Knysna lignites

(Thiergart et al. 1963). Milfordia hypolaenoides is common at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 53

Milfordia sp.

Fig. 12N

Compare

Restioniidites homeopunctatus Hekel, 1972: 15, pl. 6 (fig. 30).

Restioniidites pascuali Archangelsky, 1973: 385, pl. 9 (figs 4-8).

Description

The grain is large, ellipsoidal and monoporate. The pore is relatively small

and circular and the margin neatly defined. The exine is of medium thickness,

finely scrobiculate, and the outline of the grain is smooth. The dimensions of the

single illustrated grain are 53 x 39 wy, the pore is 8 uw wide, the exine 2 uw thick.

Discussion

Milfordia sp. is distinguished from M. hypolaenoides by its aperture

morphology, smoother exine and ellipsoidal shape. The present form is similar to

the ‘Restio subverticillatus’-type discussed in Muller (1981: 105). Since only one

specimen was observed it was not assigned to either Restioniidites homeopuncta-

tus or R. pascuali, with which it is compared.

Affinity

The affinity of Milfordia sp. is thought to be with some southern African

Restionaceae, especially ‘Restio subverticillatus’ (see discussion on M. hypo-

laenoides).

Distribution

Pollen of the ‘Restio subverticillatus’-type is known from the Maastrichtian of

north Africa (Jardiné & Magloire 1963) and the Palaeocene of south and north

America and Europe (Muller 1981). In Australia it is known from lower Eocene

to Miocene sediments (Hekel 1972; Stover & Partridge 1973). Only one specimen

was observed at Banke.

Dicolpate pollen

Genus Dicolpopollis Pflanzl, 1956

Dicolpopollis sp.

Fig. 13A—D

Description

The grain is dicolpate (disulcate?) and ellipsoidal. The colpi are three-

quarters the length of the grain and have distinct margo. The colpi are usually on

opposite sides of the grain (Fig 13C—D) but are occasionally on the same face

(Fig. 13A—B). Crassisexinous. The reticulum is fine and regular and the lumina

are circular. Length 37-43 w, width 22-30 w, lumina approximately 0,5 yp.

54 ANNALS OF THE SOUTH AFRICAN MUSEUM

oN:

Fig. 13. A-D. Dicolpopollis sp. E-I. Triorites operculatus sp. nov. J—M. Triorites sphericus

sp. nov. N. Triorites harrissii. O-P. Proteacidites sp. A. Q-R. Proteacidites sp. B.

PALYNOLOGY OF THE ARNOT PIPE 35)

Affinity

Disulcate pollen is found in extant members of the Amaryllidoideae of the

Amaryllidaceae, in some genera of the Iridaceae, in the Tofieldieae of the

Liliaceae, and in the Palmae (Chmura 1973). However, the disulcate pollen of

Monimiaceae most closely resembles Dicolpopollis sp.

Distribution

Dicolpopollis sp. is more common towards the base of the Arnot sequence.

Monocolpate pollen not assigned to genus

Forma E

Fig. 12I-J

Description

The grain is monocolpate with a relatively thin exine and all the specimens

seen had an irregular amb but were not folded. The colpus has no margo and in all

specimens was on the edge of the grain. Reticulate; the reticulum is quite coarse

and duplicolumellate. The muri and lumina are of about the same width and the

columellae usually encircle a lumina. Dimensions of illustrated specimen

5 < 26 IL:

Affinity

Forma E grains resemble the pollen of the extant palm species Areca

warburgiana (Sowumni 1972, pl. 1).

Distribution

Forma E grains are rare at Arnot.

Triaperturate pollen

Triporate pollen

Genus Triorites Cookson ex Couper, 1953

See discussion in Muller (1968: 14).

Triorites operculatus sp. nov.

Fig. 13E-I

Compare

Triorites festatus Muller, 1968: 15, pl. 3 (fig. 10).

Etymology

The specific name refers to the presence of an operculum.

56 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Triporate, spherical grains with an exine of medium width, which appears

subpsilate to granulate under the light microscope. Under the SEM it can be seen

that the exine is, in fact, psilate with minute, relatively closely spaced tuberculata

or spinules, giving the granular appearance. The pores are circular and are

surrounded by a low annulus of intermediate width, which is formed by the sexine

being slightly raised away from the nexine, i.e. forming a simple low vestibulum.

The vestibulum is so small and often inconspicuous that the grain can hardly be

described as vestibulate. The sexine may also thicken slightly in the region of the

pore. The pore has an operculum bearing the same minute tuberculata and

attached to a psilate underlying layer. The operculum is sometimes absent. The

exine is relatively thin and the grain is usually folded, this folding following no

particular pattern. Two layers are distinguishable in the exine. Diameter

. 27-30 mw, exine approximately 1 w, diameter of pore approximately 2 uw, width of

annulus 1 p.

Discussion

Triorites operculatus is different from T. festatus Muller, 1968, in having an

operculum. The pores and annulus of 7. operculatus are also smaller.

Affinity

Muller (1968: 15-16) has warned that in the case of forms that show a

generalized, primitive type of pore (as in 7. operculatus), affinity with an extant

family will be almost impossible to determine with any certainty. Broad affinity is

possibly with certain families of the Hamamelidales (Takhatajan 1969), including

the Ulmaceae, Carpinaceae, Corylaceae, Casuarinaceae and Myricaceae. Oper-

culate forms are not uncommon in these families; all the Carpinaceae (some of

which are triporate), some species of the genus Celtis of the Ulmaceae (e.g. Celtis

iguanea—Erdtman 1952), and some of the Myricaceae are operculate.

Distribution

Triorites operculatus is the dominant form at Arnot, constituting 54 per cent

in one sample, and sometimes occurring in clusters. Triorites festatus is present

from the late Cretaceous in Borneo, but is more frequent in the late Palaeocene—

Eocene and follows the same pattern in tropical west Africa (Salard-Cheboldaeff

1981). The genus Celtis is known from leaf impressions of Maastrichtian age from

the Cameroons (Salard-Cheboldaeff 1981).

Triorites sphericus sp. nov.

Fig. 13J-M

Etymology

The specific name refers to the robust spherical shape of the grain and the

fact that the grain is seldom deformed.

PALYNOLOGY OF THE ARNOT PIPE Sy

Description

Triporate grain, spherical and robust. The exine is relatively thick and

crassisexinous. The pores are circular to equatorially elongated and are

surrounded by a clear annulus. The point at which the annulus begins is marked

by a sharp line of the exine surface. The annulus is formed by the sexine

separating and rising abruptly away from the nexine. The exine is subpsilate to

faintly rugose and undulate. In some grains it appears that the exine may thicken

slightly towards the mesocolpia in the equatorial region. Equatorial diameter

20-24 w, diameter of pore opening approximately 2 w, diameter of pore and

surrounding annulus 6—7 pw, exine 2-4 w.

Discussion

Triorites sphericus differs from T. operculatus in its smaller size, relatively

thicker crassisexinous exine, lack of operculum and in sculpturing details of the

exine. Unlike T. operculatus the grain of T. sphericus is seldom folded in any way.

Affinity

See note on affinity of Triorites operculatus. Triorites sphericus 1s perhaps

morphologically closer to species of the Betulaceae or Corylaceae than to those of

the Ulmaceae, Carpinaceae or Casuarinaceae.

Distribution

Triorites sphericus is common at Arnot.

Triorites harrissit Couper, 1960

Fig. 13N

Driorites harrissii Couper, 1960: 67, pl. 12 (fig. 2). Hekel, 1972: 17, pl. 5 (fig. 7).

Description

Triporate, angulaperturate with triangular amb. Exine is relatively thick with

nexine and sexine distinct. Subpsilate, pores narrow. Equatorial diameter of

illustrated specimen 31 yw, exine 2 w, pore diameter 2 w.

Discussion

Triorites harrissii differs in shape from the two previous species. The Arnot

specimens are very similar to the illustrated specimens of Triorites harrissii of

Couper (1960) and Hekel (1972).

Affinity

The affinity of TJ. harrissii is with the Casuarinaceae or Myricaceae—more

likely the former.

58 ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Triorites harrissii has a range from the Palaeocene to the present in the

Australia-New Zealand area. Only three specimens were observed at Arnot.

Genus Triporopollenites Pflug & Thomson,

in Thomson & Pflug, 1953

There is some confusion as to what generic designation should be used for

forms with the general morphology of the species described below. The problem

arises in that four genera have been used to describe broadly similar morpho-

types. These are Proteacidites Cookson ex Couper, 1953, Triporopollenites Pflug

& Thomson, in Thomson & Pflug, 1953, Echitriporites van Hoeken-Klinkenberg,

1966, and Propylipollis Martin & Harris, 1974. A. R. H. Martin & Harris (1974)

have also discussed the problems raised by the burgeoning of the genus

_ Proteacidites and proposed two additional genera—the three being distinguished

mainly in apertural morphology.

Examples of forms that are broadly related morphologically and have been

classified under the above four genera, are the following: Proteacidites tuberculi-

formis Harris, 1965 (p. 92, pl. 29 (figs 5—7)); P. longispinosus Jardiné & Magloire,

1963 (p. 218, pl. 7 (figs 15-17)); Echitriporites trianguliformis van Hoeken-

Klinkenberg, 1966 (p. 21, pl. 42 (fig. 7)); and Triporopollenites ambiguus Stover,

in Stover & Partridge, 1973 (p. 269, pl. 21 (fig. 7)). Proteacidites tuberculiformis

Harris, 1965, was later transferred by A. R. H. Martin & Harris (1974) to

Propylipollis although, according to Harris’s original description, it lacks the

diagnostic post-atrium. Martin & Harris also excluded Proteacidites longispinosus

from Proteacidites Cookson ex Couper, 1953 (sensu Martin & Harris), without

proposing an alternative genus. To add to the confusion Boltenhagen (1978)

instituted a new species Proteacidites sigalli Boltenhagen, 1978, which he

compared to Echitriporites trianguliformis, and suggested, on very poor grounds,

that P. sigalli could be compared with pollen of the extant proteaceous genus

Spatalla.

No new genus is proposed here but it is suggested that a genus might be

considered that would include medium to large triporate forms with simple pore

structures, whose sculpturing is of scattered apicula, spinosa or micro-echina, but

which are not echinate, i.e. forms comparable, for example, to the pollen grains

of the extant proteaceous genera Telopea and Embothrium (Erdtman 1952: 356).

Triporopollenites namaquensis sp. nov.

Fig. 14A-E

Compare

Triporopollenites ambiguus Stover, in Stover & Partridge, 1973: 269, pl. 21

(fig. 7).

Proteacidites tuberculiformis Harris, 1965: 92, pl. 29 (figs 5-7).

PALYNOLOGY OF THE ARNOT PIPE

S04

Fig. 14. A-E. Triporopollenites namaquensis sp. nov. F-J. Propylipollis meyeri

sp. nov.

60 ANNALS OF THE SOUTH AFRICAN MUSEUM

Etymology

The species is named after the region Namaqualand, which in turn takes its

name from its indigenous inhabitants, the Nama.

Description

Triporate, angulaperturate pollen with a triangular amb and straight to

slightly convex sides. The grains are large and, relative to their size, thin-walled,

so that they are mostly irregular, flattened and folded. Sexine and nexine can be

distinguished and the exine is crassinexinous. The thin exine is finely punctate and

bears solitary spinules mounted on broader bases, sparsely but more or less

regularly distributed on its surface. The pollen is very variable in size and this

suggests the possible presence of more than one species. The exine thickens

slightly at the pore margins to form a distinct annular ring, which is, however,

difficult to observe in polar view. The pore is circular to equatorially elongated.

Equatorial diameter 45-90 w, pore diameter 6-10 uw, width of annulus approxi-

mately 2 uw, height of spinosa 2 pw.

Discussion

A few species of broadly similar morphology are known (see discussion on

genus). Triporopollenites namaquensis differs from Proteacidites tuberculiformis

in being spinulate (not verrucate), in the sparseness of its ornamentation, and in

having a distinct annulus—although, as stated, it is easy to miss this feature.

Except for size, the present species being considerably larger, 7. namaquensis is

very similar to T. ambiguus.

Affinity

Triporopollenites namaquensis perhaps has affinities with members of the

subfamily Grevilleoideae of the Proteaceae.

Distribution

The species is common at Arnot. Germeraad et al. (1968: 312) in discussing

the distribution of the similar morphotype, Echitriporites, stated: “The more

triangular grains with fewer and smaller spines are more common in the Upper

Cretaceous of northern South America, whereas the more rounded grains with

slightly more and larger spines are more common in the Eocene.’ The similar

Australian form 7. ambiguus is known from Palaeocene and Eocene sediments.

Genus Proteacidites Cookson ex Couper, 1953

The generic name is used here in the restricted sense as defined by A. R. H.

Martin & Harris (1974). See also discussion for Triporopollenites (p. 58).

PALYNOLOGY OF THE ARNOT PIPE 61

Proteacidites sp. A

Fig. 130-P

Description

Small, colpoidate pollen (as for example Beauprea elegans (Erdtman 1952:

343)) with triangular amb. Angulaperturate. The exine is intermediately thick,

and thins at the margins of the colpi. Sexine thinner than nexine. Reticulate.

Diameter of illustrated specimen 22 yp, exine 1,5 yp.

Proteacidites sp. B

Fig. 13Q-R

Description

Small, triporate pollen with triangular amb; a rectangle with rounded corners

in equatorial view. The pore is small and circular and the pore margins are simple.

The exine is relatively thick, crassinexinous and reticulate. Diameter of illustrated

specimen 20 w.

Affinity

Pollens similar to Proteacidites sp. B are found in the Proteaceae and the

genus Allophylus of the Sapindaceae.

Distribution

Proteacidites sp. B is rare at Arnot.

Genus Propylipollis Martin & Harris, 1974

Propylipollis meyeri sp. nov. 7

Fig. 14F-J

Etymology

This species is named for A. P. and Christine Meyer of the farm Banke.

Description

Pollen triporate, angulaperturate, amb triangular, sides straight to slightly

concave, apices roundly truncate. Exine relatively thick and crassinexinous.

Nexine thickens as the pore is approached but is truncated before the pore in the

sexine to form a post-atrium (sensu Kremp 1965, fig. 380); the state of

preservation affects the visibility of this feature. Short radiating costae pori are

present. Sexine reticulate. The reticulate pattern is irregular, angular and not

always perfect and the size of lumina decreases towards the pores, so that in the

vicinity of the pores a foveolate structure exists. Loose pieces of sexine may

sometimes be present at the pore entrance. The coarseness of the reticulum varies

62 ANNALS OF THE SOUTH AFRICAN MUSEUM

considerably and two species may be present. The muri are mostly simplicolu-

mellate, but in coarser areas duplicolumellate sections occur. The outline of the

grain is smooth. Equatorial diameter 25—33 uw, exine 2 uw, pore 2—4 w, lumina

approximately 1 w.

Affinity

The pollen of a number of genera of the subfamilies Grevilleoideae and

Persoonioideae of the Proteaceae have similar pollen. The pollen of southern

African species of Persoonioideae are not sufficiently well known to confirm a

closer affinity, but affinities to the genera Lomatia and Leucospermum have been

suggested (Germeraad et al. 1968).

Distribution

The species is common at Arnot. A related morphological form Proteacidites

dehaani Germeraad, Hopping & Muller, 1968 is common in the uppermost

Cretaceous and lowest Palaeocene strata of tropical Africa (Germeraad et al.

1968). The form is not present in late Cretaceous sediments off the south-western

Cape coast (McLachlan & Pieterse 1978) A probably conspecific form occurs in

the Neogene (?) Knysna lignites.

Propylipollis sp.

Fig. 1SA-B

Description

Triporate, the pores slightly protuberant and gaping. Angulaperturate. The

amb is triangular and the shape oblate. The exine is thick and has a complex

structure. Nexine, sexine and further subdivisions of the exine are clearly

differentiated. The sexine is thick, atectate and irregularly foveolate. The grain is

robust. Diameter of single specimen 44 w, pore 9-11 pw, exine 3-4 mw.

Only one specimen was seen and the above description should be regarded as

provisional.

Affinity

The affinity of Propylipollis sp. is perhaps with the tribe Grevilleeae of the

Grevilleoideae (Proteaceae) or with the Onagraceae.

Distribution

One specimen of Propylipollis sp. was observed at Arnot.

Tricolpate pollen

Genus Tricolpites Cookson ex Couper, 1953

See discussion by Kemp & Harris (1977: 29).

Fig. 15.

PALYNOLOGY OF THE ARNOT PIPE 63

A-B. Propylipollis sp. C-—G. Tricolpites reticulatus. H-J. Spinitricolpites

Jennerclarkei sp. nov.

64 ANNALS OF THE SOUTH AFRICAN MUSEUM

Tricolpites reticulatus Cookson, 1947

Fig. 15C-G

Tricolpites reticulata Cookson, 1947: 134, pl. 15 (fig. 45).

Tricolpites waiparensis Couper, 1960: 66, pl. 11 (figs 13-15).

Tricolpites reticulatus Cookson, 1947. Kemp & Harris, 1977: 30, pl. 5 (figs 1-2).

(See discussion by Kemp & Harris (1977) and Muller (1981: 67)).

Description

Tricolpate, fossaperturate; colpi short, extending about half-way to the

poles. The grain is circular in equatorial view, the amb is lobate, and the whole

surface is finely reticulate. The SEM photomicrographs illustrate the range of

variation in morphology. In Figure 15F the amb 1s distinctly more lobate and

fossaperturate, and the reticulate sculpture tends towards a tectate and foveolate

state; despite a narrow size range, two species may be present. The margins of the

colpi are marked by a flat seam in the sexine and a definite extension of the nexine

beyond the sexine at that point and into the mouth of the aperture. This can be

observed under both the light microscope and SEM. The grain is robust;

equatorial diameter 19-26 w, polar diameter approximately 20 uw; exine clearly

two-layered and approximately 1 mw; lumina 0,2—0,5 w.

Affinity

Tricolpites reticulatus is thought to have affinities with the genus Gunnera.

According to the data provided by Jarzen (1980) the Banke forms are the smallest

fossil Gunnera forms yet recorded, and closest—when compared to the average

for extant pollen from certain geographical regions—to those of South America

(acetolysed grains measured).

Distribution

Tricolpites reticulatus is common at Arnot, reaching a relative abundance of

7 per cent. It is known worldwide from the middle Cretaceous to the present and

is often common. See Jarzen (1980) for a full discussion of the occurrence of

Gunnera pollen in the fossil record as well as notes on the present-day habitat

requirements and distribution of the genus. A single species Gunnera perpensa, a

semi-aquatic species, is widespread in southern Africa except in South West

Africa—Namiubia.

Tricolpites sp. A

Fig. 16A-B

Description

Tricolpate; the amb is circular and the shape prolate. The colpi are long,

thin, simple slits that stretch three-quarters of the polar axis of the grain. The

exine is relatively thick and nexine and sexine are clearly differentiated. The

sexine is columellate and tectate, and the columellae are intermediately robust.

Dimensions 30-27 jw X 25-22 mw, exine approximately 1,5 w.

PALYNOLOGY OFRAHE ARNOT PIPE

Se SNE: OOS mess BZ

Fig. 16. A-B. Tricolpites sp. A. C-E. Crototricolpites densus. F-—G. Tricolpites sp. B.

- H-I. Tricolpites sp. C. K-—M. Tricolpites sp. D.

66 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

There is no information regarding the affinity of Tricolpites sp. A.

Distribution

This species was infrequent at Arnot.

Tricolpites sp. B

Fig. 16F-G

Description

Tricolpate; the amb is triangular with convexly curved sides and angulapertu-

rate. No specimen was observed in equatorial view, but it appears that its shape is

_ biconvex. The colpi are very short, narrow slits. The exine is of intermediate

height and the sculpturing is verrucate to dispersed rugulate, producing a

negatively reticulate pattern.

Distribution

Tricolpites sp. B is rare at Arnot.

Tricolpites sp. C

Fig. 16H-I

Description

Tricolpate; the amb is triangular with convexly curved sides and angulapertu-

rate. No specimen was observed in equatorial view, but it appears that the shape

is flat and very slightly biconvex. The colpi are very short, narrow slits. The exine

is of intermediate width and granulate, and apparently thins in the immediate

vicinity of the colpi to form an almost exineless rim of regular width around the

colpi. Diameter of illustrated specimen 35 w.

Discussion

In the amb, position of the apertures and short, slit-like colpi there is some

similarity between Tricolpites sp. B and Tricolpites sp. C. Only two grains of each

were seen so that their descriptions must be regarded as provisional.

Affinity

There is a tenuous similarity between Tricolpites spp. B and C and the pollen

of certain extant members of Protea such as P. mellifera or P. grandiflora

(Erdtman 1952: 350).

Distribution

Tricolpites sp. C is rare at Arnot.

PALYNOLOGY OF THE ARNOT PIPE 67

Tricolpites sp. D

Fig. 16K-M

Description

Tricolpate, amb circular to subcircular, shape oblate to biconvex. The colpi

are of intermediate width at the equator, relatively long, and narrow to a point at

their extremities. The exine is thick, and sexine and nexine are of similar width.

The nexine forms a uniform dense layer, which broadens equatorially at the colpi

margins to form marked costae endocolpi. This crassitude is apparently only

present equatorially. The sexine is of regular width and curves over the nexine at

the mouth of the colpi. The sculpturing is complex and coarse, and consists of an

irregular granulate surface covered by micro-echinae. Equatorial diameter

43-45 pw, exine 2,5—4 wu, colpi width 2,5-4 wp.

Affinity

In form, size and the unusual surface sculpturing there is some resemblance

between Tricolpites sp. D and the genus Ferocactus of the Cactaceae; see

Ferocactus latispinus (Lozano-Garcia 1979: 310, pl. 6). However, present-day

distribution and the established fossil record of the Cactaceae make this suggested

affinity somewhat unlikely and forms similar to Tricolpites sp. D occur in a

number of other families. The pollen is not similar to that of Rhipsalis, the only

extant genus of Cactaceae that is possibly indigenous to Africa.

Distribution

Tricolpites sp. D is infrequent at Arnot.

Genus Crototricolpites Leidelmeyer, 1966

Crototrico.pites densus Salard-Cheboldaeff, 1978

Fig. 16C-E

Crototricolpites densus Salard-Cheboldaeff, 1978: 224, pl. 1 (figs 10-12).

Description

The poilen grain is tricolpate, and almost circular in polar view. The grain is

invariably flattened. The colpi are broad equatorially and have gaping, ragged

margins; verrucae are the sculptural element and are angular, mostly triangular,

in polar view. Their tops are pointed, with groups of five to six arranged in a

circular pattern, the unit of a ‘croton pattern’. In polar view the colpus extends

about one-half the diameter of the grain. The equatorial diameter is 34 uw, width

of colpi at equator approximately 10 w, width of verrucae 1-2 yw, height of

verrucae approximately 1 w.

68 ANNALS OF THE SOUTH AFRICAN MUSEUM

Discussion

The Banke specimens are similar to C. densus Salard-Cheboldaeff, 1978 and

differ from C. annemariae Leidelmeyer, 1966, in the relatively smaller verrucae

and circular ‘croton pattern’ units.

Affinity

The affinity of C. densus is with the Klaineanthus-type and perhaps, because

of the doubtful feature of ragged colpi margins, with the Adenocline-subtype of

the Crotonoideae of the Euphorbiaceae (Punt 1962).

Distribution

Crototricolpites densus is common at Arnot. The genus is known from the

lower Eocene of Guyana and the Oligocene and lower Miocene of tropical

Africa. Similar forms have not been recorded from Australia.

Genus Spinitricolpites gen. nov.

Diagnosis

Medium to large-sized, more or less spherical, tricolpate spiniferous pollen

grains. The spines are medium-sized, with pointed or rounded tops and are

sparsely and irregularly arranged on the sexine.

Type species Spinitricolpites jennerclarkei sp. nov. by original designation.

Etymology

The name refers to the spiniferous and tricolpate nature of the pollen grains.

Discussion

Apart from Tricolpites latispinosus McIntyre, 1965 (p. 207, figs 13-15) no

similar forms have been encountered in the literature. The combination of large

size, spherical shape, tricolpate state and spinate sculpturing would seem to

justify the erection of a new genus. Spinitricolpites gen. nov. is here considered to

include Spinitricolpites jennerclarkei sp. nov. and Tricolpites latispinosus

McIntyre, 1965.

Spinitricolpites jennerclarkei sp. nov.

Fig. 1SH-J

Etymology

The species is named after Mr Hugh Jenner-Clarke, an exploration geologist,

who during many years work has located numerous kimberlite pipes in the

Gamoep area.

PALYNOLOGY OF THE ARNOT PIPE 69

Description

The pollen grains are medium-sized, spherical to prolate spheroidal,

tricolpate and spiniferous. The colpi are straight slits with no margo and are about

one-half to two-thirds the polar diameter of the grains. In polar view the grain is

circular but due to flattening the colpi are open at the equator. The margins of the

colpi are not strengthened in any way and may appear frayed in polar view. The

possibility that the grains are split rather than tricolpate can probably be excluded

because of the regular positioning of the colpi in polar view and because the colpi

can be seen in equatorial view. The spines are medium sized, mostly with pointed

tips, and are sparsely and irregularly arranged on the surface of the grain; they

sometimes occur in clumps or irregular rows. The exine is clearly differentiated

into a sexine and nexine and the sexine is tectate, finely columellate and punctate.

Equatorial diameter excluding spines 39-46 w, height of spines 3-5 pw, exine

1,5-2,5 w.

Discussion

The present form differs from S. latispinosus (McIntyre, 1965) in its longer,

more sparsely distributed spines.

Affinity

There is no information regarding the affinity of S. jennerclarkei. In the large

spherical shape and spinate sculpturing there is some resemblance to certain

members of the Valerianaceae or Verbenaceae (Huang 1972).

Distribution

Spinitricolpites jennerclarkei is common at Arnot; S. latispinosus (McIntyre,

1965) is known from the Miocene of New Zealand.

Tricolporate pollen

Genus Tricolporopollenites Pflug & Thomson, in Thomson & Pflug, 1953

Tricolporopollenites grandis sp. nov.

Fig. 17K-N

Etymology

The specific name refers to the large size of the grain.

Description

The morphology of the pollen is complex. It is large, tricolporate, striate, ora

lalongate or zonorate, highly prolate, ellipsoidal or with blunt ends. The colpi are

narrow and extend almost to the poles. The exine is thick and a robust, striate

sexine is clearly differentiated from the nexine. The nexine more than doubles its

width as the lalongate endoporus is approached to form a costae endoporus, and

the sexine thins in the small apocolpium. The striae are supported by wide

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 17. A-C. Tricolporopollenites brinkiae sp. nov. D-H. Tricolporopollenites arnotiensis

sp. nov. K-—M. Tricolporopollenites grandis sp. nov. O-S. Tricolporopollenites sp. A.

PALYNOLOGY OF THE ARNOT PIPE Wil

columella and sections of muri of the same width, arranged in an irregular

reticulate pattern that obscures the striate surface (which is so clearly visible in

the purely topographic SEM photomicrograph (Fig. 17N)). There seems to be a

centre line running length-wise down the middle of the mesocolpium, with a node

on the equator around which the shallow curving pattern of the striae is centred.

The colpi form a tangent to the arc of the striae with the position of the colporus

being the point of intersection of the colpi and arc. Also, the elements of the

striae are longest in the mesocolpium and apocolpium areas and are broken up

into shorter elements in the vicinity of the colporus. Length of polar axis 45-56 pw,

exine 2—5,5 w (including costae endopori), width of costae endopori 2,5-3,5 p,

sexine approximately 2 uw, width of lalongate ora or pore zone 2-3 w.

Discussion

This is an unusual grain, which is not really comparable to any known fossil.

Affinity

In size, shape, general aperture type and the complex morphology of the

sculpturing, the grain resembles the pollen of extant members of the genus

Bauhinia of the Caesalpinaceae (Senesse 1980: 394).

Distribution

Tricolporopollenites grandis is common at Arnot.

Tricolporopollenites arnotiensis sp. nov.

Fig. 17D-H

Etymology

This species is named after the Arnot Pipe.

Description

Tricolporate, amb tending towards lobate, planaperturate grain. Subprolate

in shape. The colpi are thin and about two-thirds the polar axis of the grain. The

ora are lalongate, wider than the colpi and relatively long. Sexine and nexine are

not always clearly differentiated, but the grain is crassisexinous. The sexine is

tectate and finely columellate and there may be fine suprategillar sculpturing.

Equatorial diameter 20—26 uw, polar diameter 20-24 w, ora width 1-2 w, exine

2 p, length of ora 6-7 w.

Affinity

There is some resemblance between 7. arnotiensis and the pollen of

Anthospermum (Rubiaceae). Tricolporopollenites arnotiensis also resembles

forms of the Euphorbiaceae with the Hippomane-configuration (Punt 1962), such

as Euphorbia hypericifolia and E. heterochroma illustrated in Bonnefille &

Riollet (1980, pls 47—48).

2: ANNALS OF THE SOUTH AFRICAN MUSEUM

Distribution

Tricolporopollenites arnotiensis is rare at Arnot. The extant genus Antho-

spermum is found in Africa and Madagascar.

Tricolporopollenites brinkiae sp. nov.

Fig. 17A—C

Etymology

The species is named after Brink Scholtz.

Description

The grain is tricolporate and robust. In overall shape the amb is lobate.

_ Fossaperturate. If, however, the sexine—which is very much thickened in the

equatorial mesocolpium—1is excluded, the amb of the remainder of the grain is

that of a very rounded triangle and the grain angulaperturate. In equatorial view

the grain is oblate. The sexine and nexine are clearly differentiated and the

prominent sexine is columellate, tectate and finely reticulate. The sexine thins

abruptly in the vicinity of the colpi and then bends upwards to edge the colpi with

distinct lips. The sexine is also thin in the apocolpium and the columellae are finer

here than elsewhere. Equatorial diameter 24 uw, width of colpi 2-3 w, nexine

approximately 0,5 uw, width of sexine in the mesocolpium 3 wp.

Discussion

Tricolporopollenites brinkiae has a very distinctive morphology and as far as

is known no fossil species closely resembles it.

Affinity

This species bears some resemblance to the pollen of the southern African

genus Nenax of the family Rubiaceae and, as with 7. arnotiensis, to certain

Euphorbiaceae (cf. Euphorbia heterochroma—Bonnefille & Riollet (1980,

pls 47—48)).

Distribution

Tricolporopollenites brinkiae is common at Arnot.

Tricolporopollenites coetzeeae sp. nov.

Fig. 18I-L

Etymology

This species is named after Prof. J. A. Coetzee, who has done pioneering

work on the Tertiary palynology of southern Africa.

PALYNOLOGY OF THE ARNOT PIPE

Fig. 18. A-B. Tricolporopollenites sp. C. C-F. Tricolporopollenites sp. D.

G-H. Tricolporopollenites sp. B. I—L. Tricolporopollenites coetzeeae sp. nov.

M-O. Forma H.

74 ANNALS OF THE SOUTH AFRICAN MUSEUM

Description

Large, tricolporate and verrucate grain. The exine is relatively thin and

sexine and nexine layers are difficult to distinguish, except in the region of the

thin costae endopori where the nexine is prominent. The verrucae are spherical

and large relative to the exine. However, they are decorated with microconi that

give them an angular appearance. The colpi are narrow and almost reach the

poles of the grain. The ora are broad, lalongate and short and the costae endopori

are marked. The amb is subcircular to lobate. The grain is prolate but due to the

thin exine most specimens are deformed to some extent. Length 47—52 yu, width

30-34 w, exine 2-3 w including verrucae, width of ora 4-6 pw, length of ora

9-10 pw.

Affinity

In aperture morphology, size and general exine structure there is a similarity

_ between this species and the pollen of the genus Zimmermania (subfamily

Phyllanthoideae of the Euphorbiaceae) (Punt 1962: 29, pl. 4 (fig. 1)). However,

the prolate shape of T. coetzeeae, as well as the close packing of the verrucae and

their decoration with microconi, is a condition not approached by the pollen of

any of the seven extant species of Zimmermania. The fossil form is most similar to

the pollen of the species Z. acuminata, Z. ovata and Z. capillipes (Poole 1981).

Distribution

Tricolporopollenites coetzeeae is infrequent at Arnot. The genus Zimmer-

mania is endemic to East African montane areas, where it occurs mainly in mist

forests (Poole 1981).

Tricolporopollenites sp. A

Fig. 170-S

Description

The pollen is tricolporate, the shape prolate to pointed oval, fossaperturate

and striate, the amb is circular. The colpi are fairly deep and wide, and narrow

towards the poles. The ora are lalongate and pinched at the transection with the

colpi. There are marked costae pori. The striae are simplicolumellate. Equatorial

diameter of illustrated specimen 23 w, polar diameter 32 w.

Affinity

The affinity of Tricolporopollenites sp. A is probably with the genus Rhus of

the Anacardiaceae.

Distribution

The earliest records of pollen of the Rhus-type are from the Maastrichtian of

North America and upper Palaeocene of Europe. It has not been reported from

PALYNOLOGY OF THE ARNOT PIPE 1S

the early Tertiary of Australia and is not present in Maastrichtian sediments of

southern Africa (McLachlan & Pieterse 1978). It is present in the late Neogene of

central Africa (Sah 1967) and the Neogene (?) Knysna lignites. Members of the

Anacardiaceae are known from the Eocene of tropical Africa.

Tricolporopollenites sp. B

Fig. 18G—H

Description

Tricolporate, amb circular and shape ellipsoidal. The exine is of medium

thickness and sexine and nexine are clearly differentiated. Crassinexinous. The

sexine is tectate, finely columellate and granular. The colpi are narrow and almost

reach the poles of the grain. The ora are lalongate, slit-like and of medium length

and the costae endopori are robust. Length of illustrated specimen 27 pw, width

21 w, width of ora approximately 0,5 w, length 3 wp.

Discussion

The slit-like ora is a characteristic feature of Tricolporopollenites sp. B.

Affinity

There is no information regarding the affinity of Tricolporopollenites sp. B.

Distribution

This species was infrequent at Arnot.

Tricolporopollenites spp. C and D

Species C: Fig. 18A—B; Species D: Fig. 18C—F

Description

Both species are small and angulaperturate grains. Tricolporopollenites sp. D

has a more triangular amb than Tricolporopollenites sp. C, and both are prolate.

The surface sculpturing of the two forms also differs. The colpi are narrow and

almost reach the poles. They are deeply buried in narrow clefts shaped by the

sharp inward bends in the exine at the apices of the triangular amb. The ora is

small and protrudes into this cleft producing the characteristic ‘H-shape’ seen in

equatorial view. The exine is thick and finely columellate. Length of polar axis

18-25 w, width 13-18 w, exine approximately 2 wp.

Affinity

Ferguson (1977) has described ‘H-shaped’ aperture structures in certain

genera of the Cornaceae. These structures are formed by a pore being joined to

two lateral thinnings of the endexine that run parallel to the colpus. The fossil

types described above probably possess this structure. If the uniqueness of this

76 ANNALS OF THE SOUTH AFRICAN MUSEUM

aperture morphology is confirmed, these fossils represent the first certain, and by

far the earliest, record of cornaceous pollen to date. The particular genus

concerned is Cornus and the Cornus sanguinea-subtype (Ferguson 1977: 6, figs 31,

4f-g, 5a-g). The fossil forms are unlike the pollen of Curtisia, the extant

monotypic southern African genus of the Cornaceae. Ferguson (pers. comm.

12 July 1984) notes that similar endoapertures also occur in at least one genus of

the Rubiaceae (Lewis 1965) and in various genera of the Escalloniaceae and

Penthoraceae, and in the genus Sedum of the Crassulaceae. The occurrence of

this feature is discussed in Hideux & Ferguson (1976).

Distribution

Both species are rare at Arnot.

Pollen with more than three apertures

Genus Retistephanocolpites Leidelmeyer, 1966

See Saxena (1982) for a discussion of this genus.

Retistephanocolpites sp.

Fig. 19E-G

Description

The pollen is tetracolpate. The colpi are about two-thirds the polar axis, wide

open at the equator and their margins are entire. The exine is relatively thick,

crassisexinous and tectate. The tectum is closed and columellate, and under the

SEM it can be seen that the surface sculpturing consists of a dense mat of fine

strands. The amb is circular. Diameter 16-20 pw, exine 1,5 w.

Discussion

See Saxena (1982) for a detailed discussion of the taxonomy of Tertiary

polycolpate forms. Many of these forms are known especially from the Indian

Tertiary record. As Saxena points out, exinal thickenings and precise sculptural

details are conservative features that are of the most use in attributing generic

status to fossil forms and suggesting their affinities. The unusual surface

morphology of the present form should lead to eventual positive identification.

Affinity

No positive suggestions can be made regarding the affinity of Retistephano-

colpites sp. The pollen of Rubia (Rubiaceae) and Catostemma (Bombaceae),

amongst others, appear superficially similar to this fossil species.

Distribution

This form is rare in levels sampled at Arnot but it is the dominant angiosperm

pollen in earlier (early Palaeocene?) sediments from Namaqualand.

PALYNOLOGY OF THE ARNOT PIPE

ee ee es

Fig. 19. A-B. Grootipollis reuningii sp. nov. C-D. Crotonipollis burdwanensis.

E-G. Retistephanocolpites sp. H. Ulmipollenites sp. 1-J. Forma G. K-L. Forma F.

78 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Grootipollis Krutzsch, 1966

Grootipollis reuningti sp. nov.

Fig. 19A-B

Etymology

This species is named after Dr E. Reuning, the geologist whose interest in

the Arnot Pipe, Banke, led to the discovery and study of its fossiliferous

sediments.

Description

The pollen is spherical and always invaginated to some degree. It is

periporate and has between fourteen and twenty pores. Triangular-shaped

verrucae of medium height, with rounded tops, are arranged in circular ‘croton

_ patterns’. Five to eight verrucae, each with an apex pointing towards the middle,

form a single circle, but this pattern is disrupted where a pore is situated. The

verrucae are quite small structures so that their triangular shape may not readily

be noticed in plan view. The exine is thick. Equatorial diameter 45—50 w, exine

approximately 3, size of verrucae (plan view) 1m, diameter of pore

approximately 3 w.

Discussion

Grootipollis reuningii is twice the size of the type species of the genus,

Grootipollis cretacius (Jansonius & Hills 1976: 1191), which also has relatively

smaller and fewer pores (8-10 in G. cretacius versus 14—20 in G. reuningii). Apart

from the obvious differences in aperture morphology, the smaller verrucae and

circular arrangement distinguish G. reuningii from the two other forms displaying

the ‘croton pattern’—Crototricolpites densus (see p. 67) and Crotonipollis

burdwanensis (see p. 80), which occur at Arnot.

Affinity

Grootipollis reuningii shows affinity with genera of the Thymelaeaceae (such

as Phaleria, Passerina and Struthiola) and to the Buxaceae (Muller 1981: 48). The

former is considered the more likely.

Distribution

Grootipollis reuningii is common at Arnot. The genus and related forms are

known from the late Cretaceous of the Northern Hemisphere (Muller 1981: 48),

but have not been recorded from Australian late Cretaceous or Tertiary

sediments.

Genus Ulmipollenites Wolff, 1934

See Srivastava (1969) for discussion on this genus.

PALYNOLOGY OF THE ARNOT PIPE 79

Ulmipollenites sp.

Fig. 19H

Description

Medium-sized, 5-colpate, aspidote form. The exine is relatively thick and the

aspides, around the short narrow colpi, appear as prominent knobs. The exine

appears undifferentiated under the light microscope and the surface sculpturing is

undulating to rugose. Diameter of illustrated specimen 24 yw, exine 1,5—2 w, exine

at aspides 3 yw, colpi width 0,75 yu.

Discussion

The present form differs slightly from available descriptions of other fossil

forms placed in the genus U/mipollenites and the related genus Ul/moidipites in the

degree of narrowness of the colpus pore and the degree of thickening of the

aspides. In other respects it is similar to extant species of Ulmus such as U. glabra

(Nilsson et al. 1977: 108). However, Ulmipollenites sp. is also similar to the

species Haloragis haloragoides (Cookson & Pike, 1953), suggesting a possible

affinity to Haloragis (Haloragidaceae) (H. A. Martin 1973: 21). More specimens

will have to be studied before the correct affinity of this form can be determined.

Affinity

The affinities of Ulmipollenites sp. are thought to be with the genus U/mus of

the Ulmaceae or with members of the Haloragidaceae.

Distribution

This species is rare at Arnot. Ulmus-like pollen appears over a widespread

area including Africa, North and South America and India during the

Maastrichtian (Muller 1981: 19; Salard-Cheboldaeff 1981). In West Africa the

form has a continuous record up to the Miocene. Haloragis- or Haloragicidites-

types are known from the Eocene of Eurasia (Muller 1981) and the Miocene in

Australia (H. A. Martin 1973).

Inaperturate pollen

Genus Crotonipollis Baksi, Deb & Siddhanta, 1979

The above name is used here despite the fact that Baksi et al. (1979)

instituted their genus apparently unaware of a prior homonymous generic

diagnosis with a very different content (De Lima 1976).

80 ANNALS OF THE SOUTH AFRICAN MUSEUM

Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979

Fig. 19C-D

Crotonipollis burdwanensis Baksi, Deb & Siddhanta, 1979: 233, fig. 1.

Description

The grain is large, robust and inaperturate and the sexine bears large

regular-shaped, triangular verrucae arranged in the characteristic ‘croton

pattern’. Six triangular verrucae, each with an apex pointing towards the middle,

constitute a unit of the ‘croton pattern’. The grain is so robust that under the light

microscope and normal processing no details of the nexine can be distinguished.

Diameter 50 w.

Discussion

Baksi et al. (1979) discuss the differences between two Indian species of

Crotonipollis.

Affinity

Baksi et al. (1979) mention the similarity between Crotonipollis and the

pollen of extant species of Jatropha (Euphorbiaceae). A large number of genera

of the subfamily Crotonoideae are listed by Punt (1962) as possessing the

inaperturate ‘croton’-type pollen, so that determination of closer affinities within

this group must await further work. At least some species of Jatropha have

prominently ‘ribbed’ verrucae (Bonnefille & Riollet 1980: 69, pl. 52) and thus

clearly differ from the present forms. The genus Croton is a more likely

affinity—cf. Croton draco (Lonzano-Garcia 1979: 318, pl. 12).

Distribution

Crotonipollis burdwanensis is rare at Arnot. It also has rare and restricted

occurrence in the Eocene and Palaeocene of India.

Pollen found in obligate tetrads

Genus Ericipites Wodehouse, 1933

Ericipites sp. A

Fig. 20A—C

Description

Tetrahedral tetrads of tricolporate pollen with hamulate to weakly rugulate

sculpture. The amb of each monad is subcircular. The exine is relatively thin.

Apertures arranged according to Fischer’s rule (Erdtman 1952: 14); the colpi are

long thin slits, which broaden equatorially to enclose the ora. Ora lalongate. The

costae endopori are prominent. Colpi are three-quarters to two-thirds the length

of polar axis. Diameter of tetrad 24-35 mw, exine 1-1,5 wp.

PALYNOLOGY OF THE ARNOT PIPE

G. Dicotetradites sp.

Fig. 20. A-C. Ericipites sp. A. D-E. Ericipites sp. B. F-

H-M. Triporotetradites sphericus sp. nov. N. Dicotetradites sp.

82 ANNALS OF THE SOUTH AFRICAN MUSEUM

Ericipites sp. B

Fig. 20D-E

Description

Tetrahedral tetrads of tricolporate pollen. The colpi are thin slits and

relatively short and the ora are inconspicuous. Apertures are arranged according

to Fischer’s rule (Erdtman 1952: 14). The amb is subcircular to triangular with

broadly rounded apices. The nexine is robust and the sexine is fossulate, being

traversed by fine cracks.

Affinity

As Martin (1978) and others have pointed out, the tetrads of the Ericaceae,

Empetraceae, and Epacridaceae can generally not be distinguished. The extant

families are also not well known palynologically. Because of the distribution of

the modern families it is likely that the present forms have affinity to the

Ericaceae and, less likely, to the Epacridaceae.

Distribution

The earliest record of tetrads with Ericipites-like morphology may be from

marine middle Cretaceous sediments off North Africa (Kotova 1978). Forms

designated Ervicipites are first recorded in the Northern Hemisphere in European

Maastrichtian sediments and may have affinity to the Ericaceae or Empetraceae

(Muller 1981: 41). By the Eocene they can be a common element in assemblages

from central Europe (Muller 1981). An Ericipites form, Ericipites scabratus

(Harris, 1965), comparable to Evicipites sp. A above, is an infrequent element in

middle to late Palaeocene sediments from south-eastern Australia (Harris 1965)

and a further form, Ericipites crassiexinous, is common in middle to upper

Eocene strata. Ervicipites was not recorded in Maastrichtian sediments off

southern Africa (McLachlan & Pieterse 1978) nor, for some peculiar reason, was

it observed in the Neogene (?) Knysna lignites (Thiergart et al. 1963). It is a

component of the late Neogene record in the south-western Cape (Coetzee 1981).

Ericipites sp. A is relatively common at Arnot while Ericipites sp. B is rare.

This is the earliest record from the African subcontinent of a plant group that is a

prominent member of the Capensis Flora today.

Genus Dicotetradites Couper, 1953

Dicotetradites sp.

Fig. 20F—G, N

Compare

Dicotetradites clavatus Couper, 1953: 63, pl. 8 (fig. 125).

Paripollis ochesis Partridge, in Stover & Partridge, 1973: 274, pl. 28 (fig. 2).

PALYNOLOGY OF THE ARNOT PIPE 83

Description

The pollen occurs in obligate tetrads. It could not be ascertained whether the

monads are tricolpate or tricolporate, but the colpi are long and arranged in the

normal manner according to Fischer’s rule (Erdtman 1952: 14). The ora, if indeed

present, are inconspicuous and opposite one another. The robust clavae

contribute towards the difficulty of observing the ora. Each monad is subspherical

to subtriangular; the exine is thick and clearly differentiated into a nexine and

sexine. The nexine broadens to end in a knob at the margins of the colpus. The

sexine is decorated with robust verrucae or clavae which are angular in plan and

have rounded tops. These structural elements are most robust in the distal polar

region and become smaller in the equatorial region. The sexine is hardly, if at all,

present on the interfacial proximal face of each monad. Overall size very variable.

Diameter of figured specimen 45 yw, diameter of monads 30-33 yw, exine 6 pw in

the distal polar area, 3 yw in the equatorial area.

Discussion

Only five specimens were observed and the overall size range and variation in

sculpturing suggest that more than one species is present. Figure 20N shows a

grain tending towards the clavate-baculate condition described as Dicotetradites

clavatus Couper, 1953, while the well-preserved grain illustrated in Figure 20F—G

is very similar to Paripollis ochesis Partridge, in Stover & Partridge, 1973.

Because of the uncertainty as to whether the present form has an ora or not, the

original generic diagnosis Dicotetradites was provisionally preferred and the

revised diagnosis of Crosbie & Clowes (1980) was ignored.

Affinity

The affinity of Dicotetradites sp. is probably with the Epacridaceae, which

also have obligate tetrad forms with individual grains possessing verrucate

sculpturing that may obscure their apertures, and with the sexine confined to their

distal walls and absent from the contiguous proximal walls (Mathews 1966: 464,

469, pl. 2 (fig. 3)—cf. Epacris heteronema).

Distribution

Dicotetradites sp. is very rare at Arnot. The genus Dicotetradites is a common

form in the Eocene of New Zealand with a range from the Palaeocene to the late

Oligocene (Crosbie & Clowes 1980: 460). It is also known from the Oligocene in

south-eastern Australian sediments (Stover & Partridge 1973). Similar forms

have not been reported from the Northern Hemisphere or tropical Africa. Today

the Epacridaceae are found mainly in Australia and Tasmania, but also in South

America (Willis 1966).

84 ANNALS OF THE SOUTH AFRICAN MUSEUM

Genus Triporotetradites van Hoeken-Klinkenberg, 1964

Triporotetradites sphericus sp. nov.

Fig. 20H-M

Etymology

This specific name refers to the spherical shape of the tetrad.

Description

The structure of the grain is extraordinary. It is a tetrad with each monad so

shaped that the whole grain is spherical. It could be described as a spherical, cross

tetrad with all four monads meeting at the centre. Each monad is triporate, the

pores being circular and arranged according to Garside’s rule, i.e. three pores

grouped together at four points on the surface of the grain (Erdtman 1952: 14).

The wall of each monad is psilate and is not differentiated under the light

microscope; the pore structure in this layer consists of a low, narrow but distinct

ring on the exterior of the grain, which forms the top rim of an elongated chimney

extending into the interior of the monad. Each group of pores is situated in the

hollow beneath the sexinal (?) layer formed by the curving of three adjacent

monads away from the circumference of the tetrad. The rims of the three adjacent

pores may touch one another. The four monads are enveloped by a reticulate,

simplicolumellate, undifferentiated sexinal (?) layer to form a single inaperturate

spherical grain. The reticulum is regular and perfect and the sexine may thicken

over the groups of pores. The size of the grains is very uniform, the diameter of all

measured specimens being between 31 and 34 yw, sexine varies between 1,5 and

3,5 w, diameter of pore opening 1,5-3 pw, height of whole pore structure 4—5 p,

size of lumina approximately 1 wp.

Discussion

Triporotetradites sphericus can be distinguished from Bysmapollis emaciatus

Partridge, in Stover & Partridge, 1973 (p. 273, pl. 28 (fig. 1)) in pore and exine

structure; the latter also has pores arranged according to Garside’s rule (Erdtman

1952: 14) The inaperturate, enveloping, reticulate layer of T. sphericus is the most

distinguishing feature. The only other tetrad that has pores arranged similarly is

Ajatipollis tetraedralis (Bolkhovitina) Krutzsch, 1970. In this form, however, the

pore placement is described as free, and the pores are clearly not as closely

grouped as in the present form. Crosbie & Clowes (1980: 460, figs 4, 6) note that

tetrads of the species Dicotetradites clavatus Couper, 1953, have a granulate

sexinal layer which is continuous over the junction of individual grains.

Figure 20L—M shows a grain with some of the reticulum missing and with the

psilate nexine (?) of the individual grains, as well as their pore structure, exposed.

It is likely that a new genus will eventually have to be erected to contain this form.

PALYNOLOGY OF THE ARNOT PIPE 85

Affinity

It has been suggested that members of the genus Triporotetradites have

affinity with the extant genus Gardenia of the Rubiaceae. Triporotetradites

sphericus, however, differs widely from any known Triporotetradites.

Distribution

The present form is common at Arnot. The genus 7riporotetradites is known

from the upper Eocene of Europe and the lower Miocene of the Cameroons

(Muller 1981). Muller regards a Maastrichtian record of the genus from Nigeria

(Van Hoeken-Klinkenberg 1964) as not acceptable. Stover & Evans (1973: 58)

mention an undescribed Triporopollenites type and illustrate a ‘planar tetrad’ that

superficially resembles the present form. These forms occur in late Cretaceous

and Palaeocene sediments of the Gippsland Basin, south-eastern Australia.

Dicotyledonous pollens not assigned to genus

Forma F

Fig. 19K-L

Description

Large, thin-exined 5-colpate form with a circular amb. The colpi gape

equatorially and have characteristically rounded ends. Nexine and sexine are

differentiated and the exine is subpsilate. Diameter of illustrated specimen 34 wp.

Affinity

The affinity of Forma F is perhaps with the Labiatae.

Distribution

Forma F is rare at Banke.

Forma G

Fig. 19I-J

Description

Tricolporate, very rounded triangular, planaperturate, crassiexinous, psilate

grain. Crassiendexinous. The ektexine stops short of the colpus as the endexine

thickens to form a low costae colpus. Regarding the intectate sexine the grain is

syncolpate. The endocolpus is lalongate. Equatorial diameter 25 mw, exine 2,5 wu,

endexinous costae colpus 4 w thick.

Discussion

Forma Gis a rather unusual grain with no comparable fossil forms known. In

the very thick exine and syncolpate state there is some similarity with the

following form.

86 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

The affinity of Forma G is unknown.

Distribution

Only one well-preserved grain was observed at Arnot.

Forma H

Fig. 13M—O

Description

The grain is tricolpate and crassiexinous. A thin ektexine may be present but

can hardly be differentiated from a massive endexine. At one pole the grain is

apparently syncolpate, the exine thinning towards this pole. There is some

irregularity in the margins of the colpus and a suggestion of a hexaporate

condition with the two pores of each colpus situated non-symmetrically around

the equator. The amb is lobate and the shape is oblate but with a clear

irregularity, the exine thinning towards the syncolpate pole. Equatorial diameter

of illustrated specimen 29 yw, exine 4 w, colpus 2 wu wide, polar axis 22 p.

Discussion

Forma H is a highly unusual grain especially in its lack of symmetry.

Affinity

There would seem to be little resemblance between Forma H and any extant

family except perhaps (but excluding the lack of symmetry) to the Gyrostemon-

aceae (Erdtman 1952: 198).

Distribution

Only one well-preserved grain was observed.

Forma I

Fig. 21A-B

Description

Only two specimens were observed and the following description is

preliminary. A medium-sized, probably tricolporate grain with an open reticulate

sculpturing developing into longitudinally striate sculpturing in the area of the

colpi fossae.

Discussion

As far as is known, no very similar fossil forms are known. The reticulate

sculpturing illustrated in Figure 21B resembles Alangiopollis eocaenicus as

illustrated in Reitsma (1970: 283, pl. 33).

PALYNOLOGY OF THE ARNOT PIPE

Fig. 21. A-B. Formal. C-F. Fenestriorites (photomicrographs not of material from

Arnot).

88 ANNALS OF THE SOUTH AFRICAN MUSEUM

Affinity

The affinity of Forma I is possibly with the Alangium kurzii-type of the

Alangiaceae, Section Marlea, as described by Reitsma (1970). If this is correct

then, although Forma I is a new specific record, the pattern that all early fossil

occurrences of this family are of the primitive Section Marlea is maintained.

Distribution

Forma I is rare at Arnot. Alangium kurzii and A. rotundifolium are found

today in forests of Indo-China.

POLLEN COUNTS

The pollen diagram (Fig. 3A) expresses the value of individual elements as

percentages of the total number of grains counted for each of the seven sampled

levels. Only values of one per cent and greater are included.

Little can be deduced from the pattern revealed by the diagram. Triorites

operculatus is always the most common form. A sharp increase in its relative

abundance occurs towards the top of the diagram where it achieves values greater

than 40 per cent. This increase does not correlate with simultaneous changes in

abundance of any other elements, but is preceded by a sharp increase in the

abundance of Stereisporites (Sphagnum) and the other spores. These last two

elements clearly covary. A likely explanation for this peak in their abundance is

that some change in the local geography allowed greater run-off and water trans-

port to the depositional environment. The sample from 20-21 m (65-70 feet) is

not highly carbonaceous so that the immediate presence of a peat bog is not

indicated. The increase of Stereisporites together with the other spores (which

include two possible tree-ferns— Cyathidites and a member of the Lophosori-

aceae) suggests that all grew in the same environment. This may have been a

forest with the plants concerned growing either on the floor (moss) or as part of

the understorey (tree-ferns and other ferns).

The occasional peaks in the abundance of Clavatipollenites, Triorites

arnotiensis and Tricolporopollenites brinkiae may represent changes in local

edaphic conditions and/or a slight shift in a vegetation boundary. Some extant

genera of Chloranthaceae (represented at Arnot by Clavatipollenites) are forest-

margin species. The coherent patterning in the percentage values of Triorites

operculatus and Dicolpopollis sp. are possibly indicative of long-term change in

the regional vegetation and there is some basis for speculation on its nature (see

point 21 of the discussion).

The percentage values for conifers varied between 15 and 5 per cent.

DISCUSSION

The body of this research has consisted of systematic, descriptive palynology.

The affinity of as many fossil forms as was possible has been noted and the results

of this work are summarized in point form below. The detail achieved allows for

PALYNOLOGY OF THE ARNOT PIPE 89

some comparisons to be drawn with the Australian and tropical African early

Tertiary palynomorph records, and some statistics to be produced on the

taxonomic levels and degree of extinction that has occurred in Africa between the

early Tertiary and the present.

The described assemblages from Arnot are isolated in time and space.

Although subsequent work has provided more biostratigraphic evidence, long

continuous sequences and a stable dating framework are not yet available. The

fact that only seven samples from a short 25 m sequence could be analysed makes

it difficult to evaluate the changes recorded.

The small amount of work done on palynomorph assemblages from

kimberlite pipes from the northern Cape and from Botswana and the general

paucity of published work on local late Cretaceous and Tertiary palynology make

discussion on vegetation history in these time ranges on the subcontinent

premature. However, the evidence produced in this study provides a much better

base than was previously available for tentative statements regarding the

vegetation represented at Arnot and the palaeoclimate involved. Furthermore,

since other authors (Axelrod & Raven 1978; Tankard & Rogers 1978) have

placed their interpretations upon the previously published palaeobotanical

evidence from Arnot, it was thought that some remarks on these subjects were

necessary.

Some of these remarks rely upon suggested features of the local and regional

geography of the site and its mechanism as a pollen trap for their support. These

features (see p. 5) may be summarized as follows:

The regional topography was that of a relatively mature, flat landscape

unassociated with any prominent montane region. The country rock of the area is

Namaqualand gneiss, which was blanketed by base rich ‘kimberlitic’ material in

the vicinity of the Arnot Pipe. The cones of ejectamenta of the numerous

‘kimberlite’ volcanoes are thought to have been relatively low (perhaps in the

order of 100—150 m) and the infilling of each crater to have been completed a few

million years (maximum) after emplacement. The crater lake formed in the vent

of the palaeo-Arnot Pipe volcano accumulated laminated, fine-grained, carbon-

aceous shale and mudstone sediments towards its centre and acted as a local,

small basin pollen trap, unrelated to developed drainage patterns. The agents of

pollen transport would have been wind, local run-off, and probably settling out of

suspension of fine sediments and pollen in the quiet centre of the lake after

gravity avalanching of material off the talus slope of the cone.

Although the slopes of the volcanic cones must have provided special

edaphic conditions favouring certain plant species, these areas were isolated

features in the general landscape. Even the plants that occurred on the local

edaphic site would have formed part of a more generalized, wider distribution of

plant associations. There is no climatological reason to suppose that the Palaeo-

cene location of the region experienced a transition zone between climatic regimes.

In terms of the present knowledge available, therefore, it seems safe to

suggest that the species of plants recorded at Arnot grew on base rich soils and

90 ANNALS OF THE SOUTH AFRICAN MUSEUM

were part of a widespread non-montane Palaeocene vegetation growing in the

interior of the subcontinent.

1. The following plant familes are represented in the Arnot palynoflora:

(a) Pteridophyta and Bryophyta: Sphagnaceae, Cyatheaceae, Anthocerotaceae,

Polypodiaceae, Schizaeaceae and Ophioglossaceae.

(b) Conifers: Podocarpaceae and Araucariaceae.

moideae) or Haloragidaceae (?), Casuarinaceae or Myricaceae, Proteaceae,

Gunneraceae, Euphorbiaceae, Thymelaeaceae, Anacardiaceae, Cornaceae, Eri-

caceae, Epacridaceae and Caesalpinaceae.

Several more tentative suggestions about the possible affinity of fossil

morphotypes to modern families are also made in the text.

2. The lack of diversity in the families Proteaceae, Ericaceae and Res-

‘tionaceae is notable. These three families are at present large and prominent

components of Cape fynbos vegetation. By Eocene times the Proteaeceae are

highly diversified in Australia (Martin 1981) and are represented by many forms

in the Knysna lignites (Thiergart et al. 1963), which may be either Eocene—

Oligocene (Thiergart et al. 1963; Helgren & Butzer 1977), or early Neogene in

age. Both subfamilies of the Proteaceae, the Persoonioideae and Grevilleoideae,

may be represented in the Arnot assemblages but no proteaceous form is

common to the Arnot and Australian early Tertiary assemblages. Most of the

proteaceous forms probably represent extinct genera, but the genera Leucosper-

mum and Protea may be represented. The evidence from the pollen morphology

of the Restionaceae suggests that present ideas about evolution within this family

may need revision (see Johnson & Briggs 1981).

3. The diversity in the Euphorbiaceae (non-heathland types, Specht 1981:

790) is notable. The evidence for early presence in the African subcontinent of

species with affinity to the Euphorbiaceae, Thymelaeaceae, Monimiaceae,

Anacardiaceae, Rubiaceae (?) and Ulmaceae (?), amongst others, is also

important and provides valuable data for a perspective on the in situ evolution of

plant phylogenies in the subcontinent. Wood of Euphorbiaceae and Monimiaceae

has been identified in late Cretaceous deposits on the east coast of southern

Africa (Miiller-Stoll & Madel 1962).

4. In terms of biostratigraphic age-bracketing, none of the evidence

contradicts a possible Palaeocene date for the sequence. The assemblage is

composed of a mixture of: (a) forms whose affinity to modern families can be

traced, and (b) archaic forms known mainly from, or with a record extending

back into, the Cretaceous. This combination suggests proximity to the Cre-

taceous—Tertiary boundary (see points 5 and 6 below). On the other hand

Monoporites annulatus (Gramineae) while prominent in tropical African Eocene

assemblages (Salard-Cheboldaeff 1979, 1981) does not occur at Arnot. This may

support a pre-Eocene age for these assemblages.

PALYNOLOGY OF THE ARNOT PIPE 91

5. The following form-genera and species present at Arnot are known

mainly from the Cretaceous:

Araucariacites sp., Zonalapollenites, Monocolpopollenites, Fenestriorites, Ali-

sporites grandis, Foveotriletes margaritae, Cicatricososporites, Distaverrusporites

and Hamulatisporis.

In addition Arecipites, Liliacidites, Tricolpites reticulatus and Clavatipollen-

ites forms are common in the late Cretaceous but continue through the Tertiary,

and Araucariacites australis exits from the tropical African record in the late

Cretaceous and from the Indian record at the Cretaceous—Tertiary boundary, but

has a continuous record into the Tertiary in Australia.

6. Comparing the Arnot palynomorph assemblage with other Palaeocene

assemblages* and with extant floras produces the following rough estimates:

(a) 57 per cent of the species are unique to the Palaeocene of the African

subcontinent;

(6) 59 per cent no longer occur in Africa;

(d) 39 per cent of the forms, and

(e) 60 per cent of the families, are common to the Arnot and south-eastern

Australian Palaeocene, while only

(f) 12 per cent of the forms, but

(g) 65 per cent of the families, are common to the Arnot and tropical African

Palaeocene.

(a—d excluding pteridophytes, no families of which are known to have become

extinct during the Cenozoic.)

These statistics are employed in the discussion that follows.

7. At generic levels (refer to 6e above) the Arnot Palaeocene flora is more

closely related to south-eastern Australian than to tropical African Palaeocene

floras. This reflects in the main a common southern Gondwana pteridophyte and

conifer floral inheritance. However, a degree of similarity in the climates of the

two regions is also indicated. The low level of commonality at generic rank (6a, f)

between the Arnot and tropical African assemblage suggests a marked difference

in the climates experienced in the two regions. This pattern is superimposed upon

a continuing phytogeographical relationship expressed at a higher taxonomic

level (see 6g). The uniqueness (see 6a) of the Arnot flora mainly emphasizes its

difference from the tropical African Palaeocene flora.

Points 6b and 6c are measures of the antiquity of Palaeocene floras and

although comparable estimates are not available, a rough comparison does

suggest that the level of ‘modernity’ encountered in south-eastern Australian

Palaeocene flora might be similar to that of the Arnot flora and dissimilar to the

state of the tropical African Palaeocene flora. It would appear from the work of

Salard-Cheboldaeff (1978, 1979, 1981) that this latter flora contains many archaic

* The studies on which these comparisons are based include Germeraad et al. (1968), Salard-

Cheboldaeff (1978, 1979, 1981), Martin (1978, 1981), Harris (1965), Kemp & Harris (1977),

Kemp (1981), Stover & Partridge (1973), and Muller (1981).

92 ANNALS OF THE SOUTH AFRICAN MUSEUM

forms and that, in the equatorial region, continuity with Neogene floras is only

marked from the Upper Eocene—Oligocene (Salard-Cheboldaeff 1981: 435).

The difference of 20 per cent between 6b and 6c is a measure of the level of

extinction that has occurred in Africa between the Palaeocene and the present. It

suggests the contrast in this continent between the Palaeocene environment—

with perhaps more equable climates and easy transitions between climates—and

the subsequent global development of more distinct zones of climate and

vegetation, some of which were not well represented in Africa. In particular,

Africa does not extend into high southern latitudes and therefore lacks an

extensive zone of temperate climate and the role that this might play on a

continental scale in the evolution of vegetation. For this reason, in addition to the

probable relative aridity of Africa in the Cenozoic, the subtropical—temperate

vegetation of Africa has suffered a relatively high degree of extinction between

the Palaeocene and the present.

8. The connections and contrasts of the Arnot palynoflora with early

Tertiary south-eastern Australian floras can be further elucidated.

(a) The low percentage representation of spores at Arnot relative to the

Australian norm (5% versus 20-30%) may be explained either by reference to

the general pattern of low spore representation in Africa (Salard-Cheboldaeff

1981) or by the peculiar geographic setting of the Arnot site. Relevant to the

former explanation are the suggestions that Africa has been relatively drier than

Australia for a long time and/or that the pattern reflects the intracontinental

location of the sites producing this pattern, i.e. east coast of Australia, west coast

and interior of Africa.

The latter explanation suggests that whereas it can be assumed that most

spores reach a depositional environment via water transport, this agent of

transport (as already indicated, see p. 88) may have played a minor role in the

geographical setting of the Arnot Pipe. Although involving an obviously circular

argument, there is some support in the covariation of values for Stereisporites and

the other spores for the suggestion that an increase in water transport explains the

single anomalously high value of 20 per cent recorded for these two elements at

the 65-70 foot (20-21 m) level (see Fig. 3).

(b) Apart from relatively high spore representation (indicating high humidity

and/or equability), the south-eastern Australian conifer flora is dominated by

podocarpaceous taxa, especially the genera Dacrydium and Microcachrys. In

contrast Araucariaceae entirely dominate earlier Palaeocene or late Cretaceous

assemblages from the Arnot region and make up about half of the conifer

representation at Arnot. This contrast is explained by the comparatively high

latitude of south-eastern Australia in the Palaeocene (65°S versus 40°S today) and

the correspondingly lower temperatures and more temperate climate experienced

there.

However, Araucariacites is hardly represented at all at a more northerly and

inland Australian site situated at about 50°S in the same time range (Wopfner

et al. 1974).

PALYNOLOGY OF THE ARNOT PIPE 93

Although the non-occurrence of a form at a particular site is not a reliable

observation, it is to be noted in comparing the Palaeocene Australian conifer flora

with the Arnot flora that Microcachryidites, Dacrycarpus, Dacrydium franklinii-

type, and Phyllocladus are not recorded at Arnot. All of these forms, however,

except the Dacrydium franklinii-type and the Dacrydium cupressinum-type

(recorded at Arnot) are known to occur in the late Cretaceous in the south-

western Cape (McLachlan & Pieterse 1978). Their non-occurrence at Arnot may

indicate the existence of a marked climatic gradient between the palaeolatitudes

of Arnot and the south-western Cape or it may simply indicate the existence of a

montane habitat in the latter region. It is clear, however, from evidence from

elsewhere (Herngreen & Chlonova 1981: 506, 511) that Microcachryidites is a

sensitive indicator of some climatic gradient.

Microcachryidites is still a prominent component of the Neogene vegetation

of the south-western Cape, while Araucariacites does not appear in the Neogene

record (Coetzee 1978a, 1978b, and pers. comm.). No species of Microcachrys

occur today in Africa.

It would appear that one Zonalapollenites form as well as Podocarpidites

riembreekensis and perhaps Podocarpidites kamiesbergensis are at present unique

to the southern African Palaeocene.

Australian Palaeocene from the Arnot assemblages provides further contrasts.

These include the three Nothofagus pollen types (although these are only

common from the Eocene in Australia), Myrtaceae, Olacaceae (Anacalosa),

Euphorbiaceae (Austrobuxus—Dissiliaria), Banksieae, Xylomelum-type and

other extinct forms attributed to the Proteaceae, Santalaceae, /lex (Aquifoli-

aceae) and Cupanieae (Sapindaceae).

On the other hand the following forms that occur at Arnot are not known

from the Australian Palaeocene: Triorites operculatus and T. sphericus (the

former very prominent), Retistephanocolpites (prominent earlier (?) in the

Palaeocene from other sites in Namaqualand), Crototricolpites and Crotonipollis

(Euphorbiaceae), Grootipollis (Thymelaeaceae), all the proteaceous forms

(3-4?), Milfordia (Restionaceae), Rhus (Anacardiaceae), Triporotetradites,

two monocolpate forms (one, and perhaps both, of which have affinity to

the Palmae), a palmaceous Liliacidites, Tricolporopollenites arnotiensis and

T. brinkiae (Rubiaceae, Euphorbiaceae?), and Tricolporopollenites spp. C & D

(Cornaceae).

9. Apart from the absence in the tropical African late Cretaceous—early

Tertiary record of the spore and conifer ‘temperate’ southern Gondwana floral

component, the most important contrast between the Arnot and tropical

palaeofioras lies in the representation of palms. In the Palaeocene of tropical

Africa, twelve palm form-genera are recognized and their representation is

constantly high (20-25%) (Salard-Cheboldaeff 1981). At Arnot two to three

forms with affinity to the Palmae occur and their contribution to the palynomorph

assemblages never rises above 3 per cent.

94 ANNALS OF THE SOUTH AFRICAN MUSEUM

As already indicated very few form-genera occur both at Arnot and in

Palaeocene tropical African palynomorph assemblages (12 %), while the relation-

ship is closer at a high taxonomic level, with 65 per cent of the families being

common to both.

The forms common to both regions are Foveotriletes margaritae, Distaverru-

sporis, Zonalapollenites sp. B (Cingulatipollenites), Monocolpopollenites sp. B,

Celtidoideae? (Triorites operculatus—Triorites festatus/tenuiexinus), Crototricol-

pites, Ulmipollenites sp., Proteaceae (Propylipollis meyeri—Proteacidites de-

haanii).

The families Olacaceae, Ctenolophonaceae, Malphigiaceae, Moraceae,

Acanthaceae, Mimosaceae, Bombaceae, Apocynaceae, Balanophoraceae, Mela-

stomaceae, and Combretaceae are recorded in tropical African assemblages (with

some genera being prominent) but are not known from Arnot.

A less biased reflection of differences between the tropical African and

- Arnot Palaeocene floras is provided by the list of families that, as far as is known,

occur only in the latter region: Anthocerotaceae, Sphagnaceae, Podocarpaceae,

Araucariaceae, Chloranthaceae, Myricaceae, Ericaceae, Epacridaceae, Gunner-

aceae and Cornaceae.

10. These comparisons introduce an attempt to reconstruct the palaeo-

vegetation and climate of the Arnot region. Two informal methods are used to

achieve this end. The first (points 15-18 below) involves bracketing the Arnot

climate between what has been suggested were the conditions pertaining to

palaeolatitudes to the north, south, and perhaps east of it. The second (points

18-19 below) involves extrapolating from knowledge of the present-day

distribution, habitat requirements and common growth forms of a set of taxa, to

form a hopefully coherent picture of the palaeovegetation. Table 2 provides a list

of the families and genera thought to be represented in the Arnot palynoflora,

together with notes on their present distribution, habitat and common growth

form.

11. The limitations of the second method are well known. It is clear, merely

from the presence of both Araucariacites and the Dacrydium cupressinum pollen

type at Arnot (compare the present-day habitat requirements of the genera given

in Table 2), that a process of differentiation of climates and vegetation has

probably occurred between the Palaeocene and today. Taxa such as Araucaria

and Dacrydium must have evolved relatively narrower habitat tolerances through

time, i.e. are confined to those habitats most suited to their evolutionary

potential, as a response to the development of a greater range and distinctiveness

of climates through time (cf. Kemp 1981: 40). The suggestion is therefore that

both the vegetation and climate recorded at Arnot were part of more uniform,

equable and extensive distributions of climate and vegetation (see point 13

below).

In this connection it is relevant to note that Truswell & Harris (1982: 71) in

their review of the palynology of the Eocene of Australia also mention the

occurrence of palynological assemblages containing a mixture of elements, which

PALYNOLOGY OF THE ARNOT PIPE 95

are not found growing together today, e.g. tropical rain forest and temperate rain

forest taxa.

12. Along with this observation goes the realization that it is only in a

qualified sense that concepts such as ‘tropical’ or ‘warm temperate’ can, by

extrapolation from the present, be used to describe the Palaeocene vegetation or

climates. This is not only because Palaeocene climates may have been very

different from modern climates, but also because these concepts are not basic

enough categories with which to understand even modern-day vegetation of

climatic types and distributions. Webb & Tracy (1981), for instance, use twenty-

one categories to describe the range of Australian rain-forest structural types and

nine categories to link these structural types to climatic and edaphic factors. They

argue that the recognition of the structural types enable better phytogeographical

and historical biogeographical analyses to be made between Australian and

extra-Australian regions, a claim that suggests that they have developed basic

categories with which to understand their phenomena.

13. At a general level, the mechanics of global air circulation systems are

relatively simple and therefore amenable to extrapolation to the past. Using these

mechanics and assuming the equator—pole temperature gradient of a polar-

ice-cap-free world, Lamb (1972) has modelled pre-Oligocene atmospheric

circulation. Between the equator and 60° latitude the development of pressure

anomalies would have been counteracted, not by prevailing lower and upper

atmosphere winds as is the case today, but by moving cells of higher and lower

pressures, cooler and warmer air. A weak and erratic Ferrel-type circulation

would have resulted. This type of circulation, involving the lack of zonal wind

systems, is essentially unknown today and an implication of this model is that

there are no close present-day analogues for pre-Oligocene climates and

vegetation.

This is not to suggest that, in terms of this model, some climatic gradient

between 0 and 60° latitude did not exist. An insolation gradient alone must

presumably have existed, but Lamb’s model suggests that pre-Oligocene climates

in low and middle latitudes were more uniform, perhaps more equable, extensive

and mildly transitional than is the case today. The climate of a particular region

would still have been determined by its latitude and location in terms of the

configuration of warmer and cooler seas and continents, mountains and lowlands,

and its continentality.

However, Parrish & Curtis (1982) and Parrish et al. (1982) do not accept

Lamb’s model for pre-Oligocene climates and, using an analysis of the

distribution of upwelling and organic-rich rocks and evaporites, have determined

atmospheric circulation and precipitation patterns. From this they have produced

a series of global maps portraying the broad relative isohyets of precipitation

predicted by their climatic model for various stages in the past. For the time range

concerned Parrish et al. (1982: 80) suggest that the Arnot region would have

received a relatively low to moderately low rainfall.

96 ANNALS OF THE SOUTH AFRICAN MUSEUM

Visualizing the Arnot palaeovegetation and climate in relation to those to the

north and south should be done with Lamb’s (1972) model and the predictions of

Parrish et al. (1982) in mind. It is at present not clear how real the contradictions

between the two models involved are. Kemp (1978, 1981) has used Lamb’s model

to reconstruct early Tertiary Australian climates and vegetation.

14. Both the predictions of Parrish et al. (1982) and the features of

atmospheric circulation described by Lamb (1972) allow one to maintain (contra

Axelrod & Raven 1978) that there is no immediate climatological reason why,

during the time range concerned, a zone of sharply transitional climate should

have been located in the general Arnot region producing an ecotonal state of

vegetation. Also, as stated in a previous section, there is no reason to assume that

high relief existed in the Arnot region. For these reasons, and until evidence to

the contrary exists, one should attempt to reconstruct the Palaeocene Arnot

vegetation as a single unit.

15. The comparisons made between the Arnot and tropical African

palynofloras (points 6 and 9 above) highlighted the paucity of common elements.

The Arnot palaeoflora was clearly not part of a Palaeocene African ‘tropical’

flora.

However, it should be noted that Salard-Cheboldaeff (1981) has suggested

that the climate of Africa in low latitudes during the Maastrichtian was warm,

temperate and dry and that the drastic floral change recorded at the Cretaceous—

Tertiary boundary could be explained by a cooling episode in the early

Palaeocene. Many palynological studies have documented floral change at the

Cretaceous—Tertiary boundary (Muller 1980) and a Palaeocene cooling is

generally accepted to have occurred world-wide. There is disagreement,

however, as to the scale of change attributable to this factor and its relationship to

the complex global environmental changes that took place at this time (Muller

1980). Further evidence relevant to low-latitude climates in the Palaeocene comes

from Muller’s (1980) reconstruction of the latitudinal ranges of various

thermophilous taxa through the Palaeogene. This reconstruction suggests that

their ranges were most constricted in the Palaeocene, i.e. that during the

Palaeogene temperatures were lowest in the Palaeocene itself.

16. The exiting of Araucariacites from the tropical African record during the

late Cretaceous and at the Cretaceous—Tertiary boundary in India, and its

continued occurrence in the Arnot region and in Australia in the Palaeocene, can

obviously not be explained only in terms of cooler episodes in the Cretaceous and

Palaeocene (Australia and the Arnot region at mid-latitudes would at least have

had cooler winters than the equatorial regions), but must also be related to

increased precipitation or the development of more equable climates in the

tropics. The suggestion is, therefore, that in addition to a climate cooler than that

of the tropics, a climate drier or less equable than that experienced in the

Palaeocene ‘tropical zone’ is indicated by the occurrence of Avraucariacites at

Arnot.

PALYNOLOGY OF THE ARNOT PIPE 97

However, there is a possibility that factors other than climatic change could

affect the abundance of conifers relative to angiosperms in a vegetation. Doyle

et al. (1982) have recently reviewed hypotheses explaining the prominence of

gymnosperms (Cheirolepidaceae, Araucariaceae, and Podocarpaceae) in the

mid-Cretaceous tropical floras, their subsequent decline in the late Cretaceous,

and the concomitant rise to dominance of the angiosperms. Apart from the

limited evolutionary potential of conifers in the tropics, imposed by their lack of

vessels and stereotyped leaf morphology and photosynthetic ability, the use made

by their competitors—the angiosperms— of insect pollination, to produce highly

dispersed populations and high species diversity may have contributed to their

demise. “With continued diversification, the ability of angiosperms to pack more

species into a given area might have eventually led to the collapse of gymnosperm

communities by competition from many sides and dilution of populations below a

level of effective wind pollination’ (P. J. Regal, pers. comm. in Doyle et al. 1982:

86). It should be noted that, to judge from pollen morphology, a large component

of the Arnot angiosperm palaeoflora was anemophilous, perhaps leaving the

coniferous component in a better position to maintain minimum population

densities (see point 21 below).

17. The comparisons made between the Arnot palynoflora and the south-

eastern Australian record (points 6 and 8 above) suggest that the Arnot

palaeoclimate was neither as cool nor as humid or equable as the ‘temperate’

climate indicated (especially by the composition and representation of the conifer

and spore flora) for south-eastern Australia. The odd occurrence of putative

‘tropical’ indicators such as Anacalosidites or Cupanieidites in the Australian early

Tertiary (Wopfner et al. 1974: 47) may be recording the tolerance of these taxa to

low levels of insolation in combination with high equability of climate (the effect

of eastern location of sites, a warm palaeo-Pacific Ocean, and low relief?) rather

than high ‘tropical’ temperatures.

18. As might have been predicted, therefore, the comparative sandwiching

of the Arnot palaeoclimate indicates that it was a warm, moderately equable and,

relative to the ‘tropical’ norm, dryish type. Climates of this general type are found

in subtropical and warm temperate regions today. The fact that the present-day

distributions of most of the taxa listed in Table 2 are within the tropics and

subtropics therefore lends some support to the above climatic reconstruction.

The best modern analogy for the vegetation that could have grown under

such a climate might be some of the drier forest types of east Africa, or perhaps

the mixed araucarian notophyll or microphyll vine forests of north-eastern

Australia. These are described by Webb & Tracey (1981: 626, fig. 4) as moist

forest types growing under a mean annual rainfall of 700-1 200 mm.

19. An inspection of the information on growth forms in Table 2 suggests

that the vegetation in the Arnot region was forest. Trees are the most common

growth form and lianes, epiphytes, tree-ferns, forest-floor mosses and forest-

margin species are possibly also represented.

ANNALS OF THE SOUTH AFRICAN MUSEUM

TABLE 2

Families and genera that, it is suggested, occur at Arnot, with notes on their present distribution,

size and common growth forms. (Information mainly from Willis 1966.)

Taxon

Alangiaceae (?)

(2 genera, 20 species)

Anacaridaceae

(60 genera, 600 species)

Rhus (250 species)

Anthocerotaceae

_Araucariaceae

(2 genera, 38 species)

Cactaceae (?)

(50 genera, 2 000 species)

Caesalpinaceae

Bauhinia (?) (30 species)

Casuarinaceae

(2 genera, 65 species)

Casuarina (?) (45 species)

Cornaceae

(12 genera, 100 species)

Cornus

Cyatheaceae

Epacridaceae (?)

(30 genera, 400 species)

Ericaceae

(50 genera, 1 530 species)

Euphorbiaceae

Adenocline (18 species)

or Klaineanthus (1 species)

Zimmermania (?) (4 species)

Croton (?) (750 species)

Notes

Tropics. Trees and shrubs.

Chiefly tropical, but also warm temperate areas. Trees

and shrubs.

Widespread in tropical and subtropical regions. Much-

branched shrubs, or, more rarely, trees.

Mosses mainly with a circumboreal distribution, but also

in the Mediterranean region. Hygrophytic on slightly

wet sandy soils rich in loam and mostly near forests

(Boros & Jarai-Komldédi 1975).

Southern Hemisphere, except Africa. Trees. Moist sub-

tropical and tropical non-monsoonal forests. In

Argentina Araucaria araucana is dominant in the

forest of the Subantarctic floral province.

Xerophilous growth forms of the most pronounced type.

Chiefly in the drier regions of tropical America, but

also reaching British Columbia and Patagonia. In

forest regions there are several epiphytic genera. One

genus, Rhipsalis, in Africa.

Warm regions. Mostly lianas; also trees and shrubs.

Trees or shrubs, often of weeping habit.

East Africa (? native), Mascarene Islands, Australasia.

Northern and Southern hemispheres; temperate regions

and on mountains in the tropics. Trees and shrubs,

rarely herbs.

Trees. Europe, east Asia and North America.

Tree-ferns on all southern continents.

Indo-China to New Zealand, Hawaii, South America but

chiefly Australia and Tasmania. Representing the

Ericaceae of other continents. On heaths and boggy

ground. Mostly like Ericaceae in habit, usually shrubs

or small trees.

Confined to Africa, Mediterranean and Europe in two

main masses separated by the Sahara. Cosmopolitan,

usually confined to high altitudes in the tropics; also

on moors, swamps, and peaty soils. Woody; small

undershrubs to large shrubs and a few small trees.

One of the largest plant families. Cosmopolitan in

tropical, subtropical, and warm temperate regions.

Trees in the tropics; also herbs and shrubs.

Southern Africa. Herbs.

Tropical west Africa. Giant forest trees.

Tropical east Africa. Trees.

Tropics and subtropics. Trees.

PALYNOLOGY OF THE ARNOT PIPE 99

Taxon

Gunneraceae

(1 genus, 50 species)

Lophosoriaceae

(2 genera, 2 species)

Monimiaceae

(20 genera, 150 species)

Ophioglossaceae

(4 genera, 70 species)

Palmae

(217 genera, 2 500 species)

Chamaedorea (?) (100 species)

Podocarpaceae

(6 genera, 125 species)

Dacrydium (25 species)

Polypodiaceae

(50 genera)

Proteaceae

(62 genera, 1 050 species)

Restionaceae

(28 genera, 320 species)

Rubiaceae

(500 genera, 6 000 species)

Anthospermum (?) (50 species)

Schizaeaceae

Schizaea (30 species)

Sphagnaceae

(1 genus, many species)

Thymelaeaceae

(50 genera, 500 species)

Notes

In the tropics and southern temperate regions. Perennial

thizomous herbs. Widespread in southern Africa,

except South West Africa—Namibia.

Small tree-ferns of tropical South America.

Chiefly southern tropical and especially in the ‘oceanic’

floral regions. Shrubs and trees with leathery ever-

green leaves.

Tropical and temperate regions. Small herbs, and some

tropical species are epiphytic.

Tropical and subtropical. Some are widespread, but most

genera are well localized. The palms form a charac-

teristic feature of tropical vegetation. Trees.

Warm America. Small reedy palms often forming suckers.

Southern conifers. Present on all southern land-masses.

Wide range of habitats; lowland heaths and scrubs,

open forest, rain-forest and subalpine vegetation.

Trees or shrubs.

Indo-Malaysia, Tasmania, New Zealand. Trees and

shrubs. Temperate rain-forest. Cool wet sclerophyll

forest. Mainly found in cool temperate Tasmania.

‘Frees:

Cosmopolitan, especially in the wet tropics. Almost all are

epiphytes.

Tropical Asia, Australasia, South America; tropics and

temperate areas, also mountains in Africa, South

Africa and Madagascar. The great majority live in

regions where there is annually a long dry season.

The primitive members of this family are mostly

rain-forest trees.

Mostly in southern Africa and Australia; a few in New

Zealand, Chile, Indo-China and tropical Africa.

Xerophilous, perennial with a tufted or creeping

rootstock.

One of the largest plant families. Most are tropical but a

number are temperate. Trees, shrubs and herbs.

Africa and Madagascar. Widespread, associated with

afromontane vegetation. Low shrubs and herbs.

Mainly in the tropics, but also North America. All

southern continents.

Peat-moss family. Forms peat bogs and is common on wet

forest floors and shaded mountain seeps.

Temperate and tropical regions, especially in Africa. Most

are shrubs, but there are some trees and a few lianas

and herbs.

100 ANNALS OF THE SOUTH AFRICAN MUSEUM

Taxon Notes

Ulmaceae (?) Cosmopolitan. Mainly in the Northern Hemisphere tem-

(15 genera, 200 species) perate regions and tropics.

Celtis (?) (80 species) Cosmopolitan in tropics and temperate areas. Very

widespread in southern Africa, except South West

Africa—Namibia. Trees with a range of adaptability

and growth form.

Ulmus (?) (45 species) (Elm.) North and south temperate regions. Trees.

20. Can any suggestion be made as to the habitat of members of the

Ericaceae, Epacridaceae, Restionaceae and Proteaceae, and perhaps also

Rubiaceae and Thymelaeaceae, which may not have been, or were not, trees

within the suggested general forest environment? What implications does this

have for ideas about the origins and evolution of the Cape fynbos?

. No very positive statements can be made, but there are some suggestions that

these taxa could have been part of the understorey of a dryish open forest type.

The implication is that the origin and evolution of the fynbos is linked, in its

earlier stages, to the history of this vegetation type rather than to the history of

cooler, wetter, perhaps more closed-canopy forest types, such as may have been

present to the south of Arnot, or in montane situations (Coetzee et al. 1983).

The following observations provide the basis for these statements:

(a) The counter-suggestion that the distribution of these taxa may have been

controlled by the occurrence of specific edaphic conditions receives little support

from the available evidence. There is no floristic indication of the existence of

swampy or water-logged conditions in the vicinity of the site and the suggested

local topography (p. 5) argues the same. Also, the local substrates and sediments

of the pipe itself indicate that no oligotrophic soils occurred in the vicinity of the

pipe. Therefore, Specht’s (1979, 1981) hypothesis as to the possible origin of

sclerophyllous taxa cannot, in this instance, be supported. In terms of his

hypothesis, under a warm, humid, equable climate local edaphic sites such as

water-logged areas, or areas of oligotrophic soils, could have been the areas

where early sclerophyll communities originated.

(b) Ericaceae are not recorded in the Knysna lignites (Thiergart et al. 1963),

which may be either Eocene—Oligocene or early Miocene in age, and are very

rare in the early Miocene lower levels of the Noordhoek occurrence (Coetzee

1978a, 1978b). There is evidence to suggest that the vegetation types represented

at these sites were adapted to relatively wetter and more montane and equable

climates than the Arnot palaeovegetation. Restionaceae are also either very rare

or absent from the lower levels at Noordhoek, but are the most abundant type in

the Knysna lignites. These latter restionaceous forms, however, are different to

the Arnot (Milfordia) forms, having graminoid-type apertures, and their

abundance has been taken to indicate extensive marshlands in the vicinity

(Thiergart et al. 1963).

PALYNOLOGY OF THE ARNOT PIPE 101

1983) suggests that some zonation of forest vegetation existed during the late

Cretaceous and early Tertiary in the subcontinent, and that forms with affinity to

Restionaceae, Ericaceae and Proteaceae did occur in the probably relatively drier

north-western interior.

(d) Monulcipollenites confossus, a restionaceous form, exits from the tropical

African record at the Cretaceous—Tertiary boundary (Salard-Cheboldaeff 1979).

This pattern may relate to the development of wetter and more equable climates

and closed-canopy forest during the Palaeocene. In this connection Whitmore

(1975, quoted in Webb & Tracey 1981: 613) has attributed the poverty of the

south-east Asian rain-forest grass flora to the relative stability of these humid,

closed-canopy forests. In more open and disturbable forest types (i.e. under drier

and less equable climates) the evolution of nomad grass species is favoured.

The generalized point is of importance in this discussion. Drier, less equable,

open-canopied forest types are the forest types within which an understorey will

evolve. The origins and evolution of the fynbos may therefore be linked to the

history of drier forest vegetation. In this view the fynbos shares an origin with

other generally ‘subtropical’ vegetation associations from which it has been

separated by the subsequent development of more diverse climates.

If this hypothesis is correct and it can be more adequately demonstrated that

in the early Tertiary members of the Proteaceae, Ericaceae and Restionaceae

were widespread in non-montane vegetation, then their supposedly typical

present-day distribution in Africa, high diversity in, and dominance of, the

montane vegetation of the south-western Cape, and association with nutrient-

poor soils are all ‘secondary’ features. The present pattern of their distribution

should then be viewed as being achieved as climates diversified and became, in

general (excepting especially mountains that receive orographic rain), more arid

through time. In this process earlier dominant types of vegetation and taxa were

presumably eliminated and in situ evolution of other components of vegetation

resulted in new vegetation associations, specialist adaptions, etc.

(e) The above evidence and argument suggest that Johnson & Briggs’s (1981:

463) attempt to outline the history of scleromorphic flora may need qualification.

Central to their hypotheses was the idea that, following Specht (1979, 1981), the

scleromorphic flora originated by the early Palaeogene in the adaption to patches

of oligotrophic soils within forest vegetation. Secondly, they suggest that this

scleromorphic flora has a history as a unit through into its present prominence in

areas of Mediterranean climate and poor soils.

The evidence from southern Africa suggests otherwise. In the first place, it

appears that during the early Tertiary in the Arnot region members of the

Ericaceae, Restionaceae, Proteaceae and Thymelaeaceae (and perhaps Rubi-

aceae) grew on eutrophic soils within a lowland, probably extensive, forest,

growing under a warm dryish climate. Although local edaphic factors such as

forest disturbance (fire?), steep slopes or thin soil cover may have favoured their

102 ANNALS OF THE SOUTH AFRICAN MUSEUM

growth, they must basically have been widespread within the forest, i.e. part of

the understorey. Secondly, this evidence taken together with (i) the dominance of

Restionaceae with graminoid apertures (unlike those recorded at Arnot and like

most extant southern African Restionaceae), (ii) the absence of Ericaceae in the

Knysna lignites (Thiergart et al. 1963) and (iii) the paucity of Restionaceae,

Ericaceae and Thymelaeaceae at Noordhoek (a site rimmed by mountains of

Table Mountain Sandstone, and therefore with very oligotrophic soils) in the

south-western Cape in the early Neogene (Coetzee 1978a, 1978b), suggests that

the origins of the present scleromorphic Capensis Flora are polyphyletic and that

the evolution of its components and history of its synthesis is complex.

Lastly, as suggested by Axelrod & Raven (1978) and Parrish et al. (1982),

relatively xeric vegetation may have a long history in the African subcontinent. If

this were so, comparative studies might show that relatively more of the older

scleromorphic taxa in the region were in fact truly sclerophyllous than is the case,

- for instance, in Australia.

21. The contribution of the Arnot evidence towards a preliminary outline of

the vegetation history of the African subcontinent during the late Cretaceous and

early Tertiary is discussed elsewhere (Coetzee et al. 1983). It suffices here to say

that at present the Arnot palynoflora records the first modern angiosperm flora

known in the African subcontinent after the extinction of the late Cretaceous,

archaic angiosperm flora in which Ephedripites, Fenestriorites, Cretacaeiporites,

Hexaporotricolpites and Proteacidites forms are prominent (Scholtz & Deacon

1982). The strong representation of forms such as Triorites operculatus,

T. sphericus and T. harrissii may relate to the often-recorded increase of triporate

forms with affinity to families such as the Betulaceae, Ulmaceae, Carpinaceae or

Casuarinaceae in the Palaeocene (Chourey 1974; Srivastava 1981). This phe-

nomenon has been taken, in conjunction with other evidence, to indicate a

cooling event in the early Palaeocene. Also, it has been suggested by Whitehead

(1971), amongst others, that the appearance of relatively small, triporate, psilate

angiosperm pollen indicating secondary adaptation to anemophily, coincides with

the appearance of the deciduous habit and seasonality of precipitation.

22. Lastly, the floral changes recorded within the Arnot sequence, as well as

the very different palynomorph assemblages known from the region and from

‘kimberlite’ pipe sequences from Botswana, raises a general point about the

possible differences that will be encountered in the study of vegetation change

based on ‘kimberlite’ pipe sequences, versus studies based on the more usual

depositional site sequences.

As already suggested, the crater lakes of ‘kimberlite’ volcanoes formed small

sedimentary traps positively unrelated to developed drainage patterns. It can be

assumed that their palynomorph assemblages were the result of wind and very

local water transport. In addition, the rate of their infilling is likely to have been

rapid. They will, therefore, reflect vegetation change to much finer scale than is

the case with larger epicontinental or deep-ocean sedimentary basins. This poses

PALYNOLOGY OF THE ARNOT PIPE 103

some problems for palynological work but, because of the abundance and

distribution of these sites, creates the potential for a detailed understanding of

vegetation associations, distribution and history in the time ranges concerned.

ACKNOWLEDGEMENTS

This research was funded as a project within a Council for Scientific and

Industrial Research—Cooperative Scientific Programmes (CSIR-—CSP) project,

the Fynbos Biome Project (Palaeoecology of the Fynbos Biome Subproject). The

research was originally started as part of input into the review paper “The

comparative evolution of Mediterranean-type ecosystems’ prepared by Professor

H. J. Deacon of the Department of Archaeology, University of Stellenbosch for

the CSIR-—CSP-funded conference on Mediterranean ecosystems (MEDCON

1980). The help and advice of Professor H. J. Deacon is gratefully acknowledged.

Mrs C. E. Stevens of the Department of Archaeology, University of

Stellenbosch, gracefully typed and retyped the manuscript, and painstaking care

was taken in producing the photomicrograph prints by Elsabé Pretorius. Many

thanks also to Mr A. P. and Mrs Christine Meyer, and to Esmien and Jannie

Louw of the farms Banke and Riembreek respectively, for their friendship and

hospitality.

Helpful criticism of the manuscript was received from Professor E. J. Moll,

Dr L. Scott and Dr E. M. Truswell (formerly Kemp). Dr M. Cluver of the South

African Museum kindly made the samples available for study. I am very grateful

to Miss E. Louw for considerable editorial assistance.

The University of Stellenbosch provided a generous grant towards part of the

publication costs of this work.

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PALYNOLOGY OF THE ARNOT PIPE 109

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7 & “2

: = .

F <

= = t 7

aa i 1—& a

= 8 x ri Se =

r F a -

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)

ae i a -

= 7 : ‘

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a = SSS Se —_—_—SSSSES__S__Co ee ees = = = SSS = = ———— —— ——— —

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

(particularly Articles 22 and 51).

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

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

nov., syN. nov., etc.

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

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

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

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

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

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

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

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

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

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

Nucula largillierti Philippi, 1861: 87.

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

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

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

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

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

not acceptable.

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

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

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

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

must be recorded, e.g.:

Holotype

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

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

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

7. SPECIAL HOUSE RULES

Capital initial letters

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

>’ s

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

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

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

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Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

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

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

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

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

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

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

Biological Abstracts.

A. SCHOLTZ

THE PALYNOLOGY OF THE

UPPER LACUSTRINE SEDIMENTS

OF THE ARNOT PIPE,

BANKE, NAMAQUALAND

VOLUME 95 PART 2 APRIL ISSN 0303-2515

CAPE TOWN

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Title: informative but concise, without abbreviations and not including the names of new genera or species

Author’s(s’) name(s)

Address(es) of author(s) (institution where work was carried out)

Number of illustrations (figures, enumerated maps and tables, in this order)

(b) Abstract of not more than 200 words, intelligible to the reader without reference to the text

(d) Introduction

(e) Subject-matter of the paper, divided into sections to correspond with those given in table of contents

(f) Summary, if paper is lengthy

(g) Acknowledgements

(h) References

(i) Abbreviations, where these are numerous

3. MANUSCRIPT, to be submitted in triplicate, should be typewritten and neat, double spaced

with 2,5 cm margins all round. First lines of paragraphs should be indented. Tables and a list of

legends for illustrations should be typed separately, their positions indicated in the text. All

pages should be numbered consecutively.

Major headings of the paper are centred capitals; first subheadings are shouldered small

capitals; second subheadings are shouldered italics; third subheadings are indented, shouldered

italics. Further subdivisions should be avoided, as also enumeration (never roman numerals)

of headings and abbreviations.

Footnotes should be avoided unless they are short and essential.

Only generic and specific names should be underlined to indicate italics; all other marking

up should be left to editor and publisher.

4. ILLUSTRATIONS should be reducible to a size not exceeding 12 « 18 cm (19 cm including