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Neogene palynomorphs from the Huon Peninsula, Papua New Guinea

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DOI: 10.1080/01916122.1982.9989233 · Source: OAI

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Neogene palynomorphs from the Huon peninsula,

Papua New Guinea

Geoffrey Playford

To cite this article: Geoffrey Playford (1982) Neogene palynomorphs from the Huon peninsula,
Papua New Guinea, Palynology, 6:1, 29-54, DOI: 10.1080/01916122.1982.9989233

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 NEOGENE PALYNOMORPHS FROM THE
HUON PENINSULA, PAPUA NEW GUINEA
GEOFFREY PLAYFORD
Department of Geology and Mineralogy
University of Queensland
St. Lucia, Brisbane
Australia 4067

Abstract
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Palynological analysis of outcropping claystones and shales from
three low-grade coal occurrences in the Pindiu area, central Huon
Peninsula of Papua New Guinea, discloses well-preserved and
generally similar assemblages of spores and pollen grains. These
occurrences are from within incompletely known but dominantly
marine strata considered to range in age from early Miocene to
Pliocene. A total of 25 form-specific categories of spore-pollen
palynomorphs are recognizable in the assemblages, together with
one type ("grade") of epiphyllous fungal "germling". One new
genus, Scolocyamus, typified by S. magnus sp. nov., is
established, as are the following new species: Biretisporites huon-
ensis, Baculatisporites scabridus, Matonisporites mulleri, Conver-
rucosisporites ponderosus, Perfotricolpites maculosus, Rostria-
pollenites robustus, and Margocolporites tricuneatus.
Botanical alliances of the dispersed palynomorphs are mainly
with the ferns, among which several families, including the Poly-
podiaceae and Blechnaceae (extant genus Stenochlaena), are
represented. A variety of angiospermous groups is also
represented, but gymnospermous elements are minimal. Despite
the evidently close stratigraphic association with marine deposits,
the palynofloras themselves contain no obvious forms of marine
derivation. The assemblages appear to represent the autochtho-
nous products of tropical freshwater swamp vegetation of seem-
ingly low diversity.
INTRODUCTION
1976a, b) has examined the palynology of cored
sediments from an onshore oil-exploration well (Iviri
No. 1), located also in the Central Delta region. In
his 1976 papers, Khan described Late Tertiary spores
and pollen, including many new species, and fore-
shadowed "later papers (to) consider the climatic,
ecological, phytogeographical and stratigraphical
implications" (Khan, 1976a, p. 753). These had not
appeared at the time of writing, but are awaited with
interest.
Morley (1978) has produced a valuable conspectus
of the Cenozoic spore-pollen record in southeast
Asia. He emphasized, inter alia, that the complexity
of regional vegetational patterns will presumably
necessitate the establishment of separate palyno-
stratigraphic zonal schemes for the various regions.
The present paper represents the initial record of
Late Tertiary spore-pollen suites from the Huon
Peninsula of Papua New Guinea, in an area some
350 km east-northeast of the Central Delta region.

Published accounts of the palynology of Late

Tertiary sediments in Papua New Guinea are by no
means extensive, certainly in comparison with
available information on the Quaternary of that
region (see, for example, Flenley, 1979, pp. 88-94).
Cookson and Pike (1953, 1954) and Cookson (1957)
have described selected species of spores and pollen
grains from coals, said to be Pliocene in age, from
the Central Delta region around the head of the Gulf
of Papua (i.e., from the south-central margin of the
onshore Papuan Basin). More recently, Khan (1974;
Palynology, 6: 29-54 (1982)
GEOLOGICAL SETTING
The Huon Peninsula, on the central east coast of
Papua New Guinea (Text-Figure 1) occupies the east-
ernmost part of the North New Guinea Basin,
specifically of its constituent Ramu Sub-basin
(Grund, 1976). The peninsula takes in the Huon and
southernmost Sag Sag 1:250,000 Sheet areas, and has
been mapped geologically on the scale by officers of
the Geological Survey of Papua New Guinea
(Robinson, 1974a, b).
 30
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Text-Figure 1. Locality map of central part of Huon Peninsula, showing numbered localities (1-5) of Neogene coal occurrences (after Malagun
and Tamu, 1981, fig. 1). Palynomorphs were obtained from localities 1, 3, and 5.
The rugged topography, with its extensive and
dense rainforest cover and severely limited accessi-
bility, have so far inhibited a detailed understanding
of the peninsula's stratigraphy and structure, which
are undeniably complex. Robinson (1974a, b) has
differentiated some sixteen rock units, many of them
lateral equivalents, totalling ca. 10,000 m in thick-
ness. The sequence ranges in age from Oligocene to
Miocene, principally on the basis of foraminiferal
faunas, and comprises in very broad terms: a
volcanogenic sequence, up to 5500 m thick and early
Oligocene to early Miocene in age, succeeded by
mainly shallow water marine strata, dating from the
early Miocene and consisting chiefly of reefal carbo-
nates and associated clastic sediments.
The occurrence of lignite in the Huon Peninsula
was initially and briefly reported by Robinson
(1974a, b) in the Kabwum area of the peninsula's
PALYNOLOGY, VOLUME 6 — 1982
northwestern interior. This deposit is in the form of
"impure argillaceous and calcareous lignitic beds up
to 4 m thick" within the basal part of the Tipsit
Limestone (early to middle Miocene; up to 500 m
thick). In 1979, Malagun and Tamu participated in
an 'Operation Drake' reconnaissance expedition, and
in 1981 they published a brief account of lignite and
higher grade coals they had encountered at four
localities in the general area of Pindiu, central Huon
Peninsula. As discussed subsequently, samples from
three of those localities, and from an additional one,
were analyzed palynologically in the present study.
Unfortunately, due to limitations imposed by time
and difficult terrain, Malagun and Tamu were unable
to document the coal occurrences in terms of strati-
graphic or structural detail. Nevertheless, it is clear
from their account, and from the 1:250,000 geolog-
ical map (Huon-Sag Sag Sheet; Robinson, 1974a, b),
that the samples were collected from Neogene
(Miocene or Pliocene) stratal sections.
 Neogene Palynomorphs from Papua New Guinea
MATERIAL AND METHODS
The locations of all the collection sites for coal and
associated carbonaceous sediments in the Pindiu
area, central Huon Peninsula, are depicted in Text-
Figure 1. The first four numbered localities are
respectively those of the original collectors (Malagun
and Tamu, 1981, fig. 1). Locality 5 is an additional
coal occurrence of Malagun and Tamu but was not
cited in their paper. No sample was available for
palynological analysis from locality 2.
Details pertaining to the studied samples and their
locations are given below, as provided by Malagun
and Tamu (1981) and Dr. D.W. Haig (oral commun.,
June 1981; written commun., 19 October 1981).
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Locality 1. Botare Creek, tributary of Suamu
River; approximately Lat. 6° 27' S., Long.
147o34'/2' E. Dark grey-brown, carbonaceous clay-
stone; palynological preparation no. D503; with rare
benthic and planktic foraminiferids said to be of
deepwater marine origin and of Pliocene age (pos-
sibly reworked?).
Locality 3. Foria River, approximately Lat.
6°24!/2' S., Long. 140°31!/2'E. Mid-grey, friable,
carbonaceous claystone; palynological preparation
no. D504. Abundant associated foraminiferids, Cel-
lanthus craticulus (Fichtel & Moll) and Ammonia
beccarii (L.) (both benthic "smaller" rotaliines), are
known to range upwards from early and middle
Miocene respectively. "A coastal hyposaline
swamp" was suggested as possible paleoenvironment
(Haig in Malagun and Tamu, 1981, p. 53).
Locality 4. Urawa Creek, subtributary of Kua
River, approximately Lat. 6°27' S., Long.
147°28'E. Lignite, containing more than 90 per cent
of vitrite, associated with pyrite and rare sclerotia
(Dr. G. Khorasani, written commun., 12 November
1981); termed by Malagun and Tamu (1981, p. 53) "a
subbituminous to bituminous coal" interbed (less
than 3 m thick) within a coralline limestone sequence
totalling 50 m in thickness. The coal proved to be
palynologically barren.
Locality 5. Walking track on east side of Mawu
Creek, a southflowing tributary of Kua River;
approximately Lat. 6°23i/2' S., Long. 147°261/2' E.
Sample provided consists of black carbonaceous
shale, together with dull lignite containing vitrinite
and abundant pyrite and large amounts of organic
31
detrital material (inertodetrinite + liptodetrinite)
according to Dr. G. Khorasani (written commun., 12
November 1981). Palynological preparation no.
D502 (from the shale).
Approximately 5 g of each sample were processed
for the extraction and concentration of plant micro-
fossils. Briefly, the laboratory techniques entailed:
initial treatment of washed, coarsely crushed sample
with 10 per cent hydrochloric acid, for dissolution of
carbonates; boiling for 30 minutes in 33 per cent
hydrofluoric acid, for removal of mineral matter; a
single treatment with hot 50 per cent hydrochloric
acid, for dissolution of fluoride precipitates;
oxidation by 5-8 minutes' immersion in warm
concentrated nitric acid; washing with very weak (ca.
0.5 per cent) ammonium hydroxide; acetolysis (fol-
lowing Erdtman, 1960); screening through 171 /xm
nylon mesh; and heavy-liquid separation (zinc
bromide solution, S.G. 1.5). After each step, the
residues were thoroughly washed to neutrality. The
ultimate residues were stored in small, stoppered
plastic bottles in glycerol:water (1:1) mixture. Three
of the four samples, from localities 1, 3, and 5,
yielded abundant, well-preserved plant microfossils;
the remaining sample, from locality 4, proved to be
totally devoid of palynomorphs.
Several strew slides and numerous single-specimen
slides were prepared from each productive residue
for light-microscopic study, using glycerine jelly as
mounting medium. Selected spore and pollen species
were examined under the scanning electron micro-
scope (Cambridge Stereoscan I1A instrument); the
specimens thus studied were finally remounted in gly-
cerine jelly on conventional slides.
SYSTEMATIC PALEONTOLOGY
In this section, form-species described, or recorded
with comment, are arranged into broad morphologi-
cal categories above the level of form-genus. Natural
affinities are cited where known, or inferable with
varying degrees of confidence. Newly instituted
form-taxa, of which there are one new genus and
eight new speices, are each based upon at least 10
suitably preserved and oriented specimens. All illus-
trated specimens (holotypes, paratypes, and hypo-
types) are deposited in the micropaleontological type
collection of the Department of Geology & Mineral-
ogy, University of Queensland, Brisbane. Each
specimen is designated, as in ensuing text and plate
 32
explanations, according to the following sequence:
preparation/slide number, east-west and north-south
mechanical-stage readings (of Zeiss Photomicro-
scope II, no. Mx3237 of the above-cited Depart-
ment), and registered specimen number (prefixed
'Y').
Trilete Spores
Genus Biretisporites Delcourt & Sprumont
emend. Delcourt, Dettmann, & Hughes 1963
Type-species: Biretisporites potoniaei Delcourt &
Sprumont 1955; by original designation.
Biretisporites huonensis sp. nov.
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Plate 1, figures 9-13
Diagnosis: Spores radial, trilete. Amb sub-
triangular, with rounded apices and convex to almost
straight sides. Laesurae distinct, extending 0.4-0.7 of
distance to equator, often unequal on a given
specimen; accompanied (in well-preserved examples)
by very narrow, elevated lips {ca. 1 /im in overall
width). Exine laevigate to scabrate, 1.2-1.5 ,um
thick, often displaying arcuate compression folds.
Dimensions (30 specimens): Equatorial diameter
54 (63) 72 /an.
Holotype: Preparation D504/3, 89.1 20.3,
Y.356I; Plate 1, figure 11. Amb roundly subtri-
angular, diameter 61 ^m; laesurae narrowly lipped,
length ca. 0.7 of spore radius; laevigate exine 1.3 ^m
thick, with one major compression fold.
Type locality: Locality 3.
PLATE 1
1-4 Baculalisporiles scabridus sp. nov., X750. 1, Holotype,
median focus; D5O3/1, 101.0 8.3, Y.3551. 2, Median focus;
D5O3/1, 120.0 16.1, Y.3552. 3, Distal focus; D5O3/1, 93.9
21.0, Y.3553. 4, Distal focus; D5O3/1, 111.2 9.7, Y.3554.
5 Foveosporites sp. B, X500. Median focus; D502/1, 107.6
17.2, Y.3555.
6,7 Foveosporiles sp. A. 6, Distal focus, X500; D502/2, 109.7
13.2, Y.3556. 7, Proximal focus, X75O; D504/2, 121.4
13.3, Y.3557.
PALYNOLOGY, VOLUME 6 — 1982
Derivation of name: After Huon Peninsula,
Papua New Guinea.
Comparison: Biretisporites huonensis sp. nov.
differs from Cyathidites australis Couper 1953 in its
more roundly triangular amb, and shorter and
typically unequal laesurae that are, moreover,
narrowly lipped. None of Krutzsch's (1962)
numerous taxa of laevigate trilete spores from the
Tertiary of north-central Europe could be considered
as specifically comparable with B. huonensis.
Affinity: Other than presumably pterophytic, no
definite decision can be reached as to the botanical
affinity of Biretisporites huonensis.
Occurrence: Localities 1,3, and 5.
Genus Baculatisporites Pflug & Thomson
in Thomson & Pflug 1953
Type-species: Baculatisporites primarius (Wolff)
Pflug & Thomson 1953; by original designation
(monotypic).
Baculatisporites scabridus sp. nov.
Plate 1, figures 1-4
Diagnosis: Spores radial, trilete, with circular to
subcircular amb. Laesurae sinuous, extending to ca.
0.8 of distance to equator; accompanied by narrow
elevated lips that often show gradual increase in
width to radial termini where slight curvaturae may
be developed. Exine sculptured with a mixture of
bacula and spinae (typically blunted) with some ver-
rucae interspersed; sculptural elements discrete (up to
8 Rugulatisporites trophus Partridge in Stover & Partridge
1973. Proximal focus, X75O; D5O3/1, 115.4 10.9, Y.3558.
9-13 Biretisporites huonensis sp. nov. 9, Proximal focus, X500;
D504/2, 123.0 3.8, Y.3559. 10, Median focus, X500;
D504/2, 90.7 3.9, Y.3560. 11, Holotype, proximal focus,
X500; D504/3, 89.1 20.3, Y.3561. 12, Proximal surface,
X1500; D5O4/S1/13, 114.5 12.2, Y.3562. 13, Proximal sur-
face, X1500; D504/S1/5, 114.2 12.9, Y.3563.
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Neogene Palynomorphs from Papua New Guinea

PLATE 1
 34
7 jim apart) or fused basally in groups of 2 to 4; bases
circular in outline (basal diameter 0.8-3.2 jan);
length 1.8-5 [im. Sculpture most prominently
developed on whole of distal hemisphere and equa-
torially on proximal hemisphere; contact faces bear
only scattered, relatively small projections. Thick-
ness of non-apiculate exine 1.5-2.2 pun.
Dimensions (12 specimens): Equatorial diameter,
excluding sculptural projections, 43 (50) 54 pirn.
Holotype: Preparation D5O3/1, 101.0 8.3,
Y.3551; Plate 1, figure 1. Amb subcircular, 51 /xm in
diameter; exine 2.2 pun thick, bearing somewhat
heterogeneous apiculate sculpture, dominantly on
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distal surface and on equatorial margin of proximal
surface; sculptural projections mainly blunted to
acute-tipped spinae and bacula, basally coalescent or
discrete (up to 5 pun apart), length 1.9-4.5 pun, basal
diameter 1-2.5 pirn; contact areas irregularly punc-
tate (?corrosion effect) and bearing only sparse
apiculate projections, which are entirely absent in
proximal polar region.
Type locality: Locality 1.
Derivation of name: Latin, scabridus, rough.
Comparison: Baculatisporites papuanus, as de-
scribed by Khan (1976a) from the Neogene of the
Central Delta region of Papua New Guinea, is more
coarsely and uniformly sculptured, and is generally
larger than B. scabridus sp. nov. Saxosporis bran-
denburgensis Krutzsch 1967, of the European
Tertiary, is also larger than B. scabridus and
possesses finer sculpturing elements.
PLATE 2
1-7 Polypodiaceoisporites retirugatus Muller 1968; 1-6, X500;
7, X75O. 1, Proximal focus; D503/2, 121.8 9.3, Y.3564. 2,
Proximal focus; D503/2, 104.6 13.6, Y.3565. 3,Proximal
focus; D5O3/1, 88.7 14.3, Y.3566. 4, Median focus;
D5O3/1, 96.5 6.5, Y.3567. 5, Median focus; D5O3/2, 101.9
8.9, Y.3568. 6, Proximal and distal foci respectively;
D503/1, 110.1 17.3, Y.3569. 7, Proximal focus; D504/2,
94.1 20.2, Y.3570.
8-10 Matonisporites mulleri sp. nov., X750. 8, Holotype, proxi-
mal focus; D5O3/1, 91.8 13.1, Y.3571. 9, Median focus;
PALYNOLOGY, VOLUME 6 — 1982
Affinity: The morphology of this form suggests
either bryophytic (i.e., anthocerotacean: Jarzen,
1979) or fern derivation.
Occurrence: Locality 1.
Genus Converrucosisporites Potonie & Kremp 1954
Type-species: Converrucosisporites triquetrus
(Ibrahim) Potonie & Kremp 1954; by original
designation.
Converrucosisporites ponderosus sp. nov.
Plate 2, figures 13-15
Diagnosis: Spores radial, trilete; amb convexly
subtriangular. Laesurae simple, length ca. 0.6-0.8 of
spore radius; sometimes partly bordered by exinal
compression-folds. Exine 5-6.5 pun thick, bearing
verrucae that are conspicuously developed on distal
and proximo-equatorial regions, but are normally
sparse on contact areas. Verrucae may locally be
fused in pairs, but are usually discrete (0.5-6 pun
apart) with laevigate to scabrate intervening exine.
Verrucae circular or subcircular in basal outline
(diameter 1.2-4.5 ^m, rarely as much as 9 pirn),
height 1.3-4 pun, broadly rounded in lateral aspect.
Dimensions (12 specimens): Equatorial diameter
92 (102) 116 pun, sculpture excluded.
Holotype: Preparation D504/15, 104.1 10.5,
Y.3576; Plate 2, figure 13. Amb subtriangular with
convex sides and broadly rounded apices; equatorial
diameter 110 /<m; laesurae about three-fifths of spore
D5O3/1, 91.8 10.2, Y.3572. 10, Proximal focus; D5O3/1,
112.6 17.6, Y.3573.
11,12 Cf. Rugulatisporites sp., X500. 11, Proximo-equatorial
aspect; D5O3/1, 122.6 12.7, Y.3574. 12, Median focus;
D503/2, 101.6 21.2, Y.3575.
13-15 Converrucosisporiles ponderosus sp. nov.; median foci,
X500. 13, Holotype; D504/15, 104.1 10.5, Y.3576. 14,
D504/3, 98.1 12.0, Y.3577. 15, D504/2, 101.0 22.3,
Y.3578.
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Neogene Palynomorphs from Papua New Guinea

PLATE 2
 36
radius; exine 5.5 /xm thick; verrucate sculpture com-
prehensive but sparsely developed proximally in
contact areas, verrucae mostly circular in basal
outline, diameter 1.5-4 (*m, height 1.5-3 /xm.
Type locality: Locality 3.
Derivation of name: Latin, ponderosus, heavy.
Comparison: These spores differ from those of
Verrucosisporites kopukuensis (Couper) Stover (in
Stover and Partridge, 1973) by having a generally
thinner walled exine that possesses a denser and more
variable, albeit predominantly verrucate, sculpture.
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Affinity: Unknown, but possibly with the ferns.
Occurrence: Locality 3.
Genus Foveosporites Balme 1957
Foveosporites sp. A
Plate 1, figures 6, 7
Remarks: Three spores only of this species have
been observed, ranging from 43 to 51 /xm in
equatorial diameter. Their negative sculpture,
confined to the distal surface, consists of fine
irregular pits and vermiculi (the latter being more
conspicuous towards equator). The specimens show
some resemblance to Foveotriletes lacunosus
Partridge (in Stover and Partridge, 1973), but the
latter is differently sculptured in detail and is
generally smaller.
Affinity: These spores are akin to those of the
Lycopodium phlegmaria L. type (see Erdtman and
Sorsa, 1971, p. 113).
PLATE 3
1 Gleicheniidites circinidites (Cookson) Dettmann 1963.
Proximal focus, X500; D503/1, 19.9 94.3, Y.3579.
2,3 Laevigatosporites major (Cookson) Krutzsch 1959. 2,
Lateral aspect; D5O3/1, 95.5 18.2, Y.358O. 3, Proximal
focus; D502/3, 126.6 11.8, Y.3581.
4-11 Microfoveolatosporis spp. 4, Proximal focus, X500;
D502/2, 110.6 13.0, Y.3582. 5, Proximal focus, X750;
D504/2, 93.4 12.4, Y.3583. 6, Median focus, X750;
D502/3, 94.2 14.8, Y.3584. 7, Laterial aspect, X500;
PALYNOLOGY, VOLUME 6 — 1982
Occurrence: Localities 3 and 5.
Foveosporites sp. B
Plate 1, figure 5
Remarks: Another rare, negatively sculptured
trilete form with subcircular amb (diameter
80-91 /xm, 3 specimens only). Laesurae, extending to
four-fifths of distance to equator, are flanked by lips
up to 3.5 /xm in overall width and height. Exine 4 /xm
thick; laevigate to faintly wrinkled in contact areas,
distinctly foveolate-vermiculate elsewhere, most
prominently on the distal hemisphere; depressions
mostly of irregular shape, disposition, and size,
0.5-6 pirn in maximum diameter or length,
0.5-1.5 /xm deep, up to 6 /xm apart (usually ca.
2.5 /xm).
Affinity: This form could be related to either the
Lycopodophyta or to the pterophyte family Ophio-
glossaceae (cf. Harris, 1955; Erdtman and Sorsa,
1971).
Occurrence: Locality 5.
Genus Rugulatisporites Pflug & Thomson
in Thomson & Pflug 1953
Rugulatisporites trophus Partridge
in Stover & Partridge 1973
Plate 1, figure 8
Remarks: The single specimen, 54 /xm in equator-
ial diameter, displays the characteristic features of
Partridge's species, as originally described from
Eocene-Oligocene strata of southeastern Australia.
The comprehensive exinal sculpture consists of
coarse, close-spaced rugulae (3-9 /xm wide) that
irregularly branch but do not anastomose to form a
D504/3, 108.8 10.8, Y.3585. 8, Lateral aspect, X500;
D503/1, 94.1 7.2, Y.3586. 9, Median focus, X750; D504/2,
122.1 20.0, Y.3587. 10, Proximo-equatorial aspect, X1000;
D504/S1/15, 112.7 11.2, Y.3588. 11, Disto-equatorial
aspect, X1750; D502/S1/20, 109.8 14.5, Y.3589.
12,13 Polypodiidites sp. 12, Distal surface, X1000; D502/S1/13,
110.2 15.6, Y.3590. 13, Proximal surface, X1500;
D502/S1/7, 111.1 15.6, Y.3591.
 Neogene Palynomorphs from Papua New Guinea PLATE 3

\
' 3

•* • I
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\ -.

if

I I

10

& &*>19miS3Ll&&m
 38
reticulate pattern. Exine is 6.5-8.5 nm thick,
including the rugulae that, moreover, tend to
coalesce adjacent to laesurae to form lip-like
structures.
Affinity: This form is possibly related to the
Lycopodophyta (cf. spores of Lycopodium clavatum
group in Knox, 1950, pi. 10).
Occurrence: Locality 1.
Cf. Rugulatisporites sp.
Plate 2, figures 11, 12
Remarks: This informal category is represented by
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two large trilete spores, 91 ^m and 105 pm in
diameter. Distinct, straight, simple laesurae, length
0.7-0.8 of spore radius. Contact faces laevigate; else-
where, exine is sculptured with low, smooth, rounded
elevations that are mainly irregular rugulae,
4.5-13 fttn broad, with some interspersed verrucae.
Unsculptured exine is 3.1-3.8 ^m thick.
No closely comparable form is evident from
available literature.
Affinity: Uncertain, but possibly with the ferns.
Occurrence: Locality 1.
Genus Matonisporites Couper
emend. Dettmann 1963
Type-species: Matonisporites phlebopteroides
Couper 1958; by original designation.
Matonisporites mulleri sp. nov.
Plate 2, figures 8-10
Diagnosis: Spores radial, trilete. Amb near-
triangular with rounded apices and almost straight
sides. Laesurae distinct, sinuous, attaining equatorial
margin; accompanied by narrow, elevated lips, about
1 ^m in overall width. Exine laevigate, thicker at
amb apices (2.8-4 ^m) than along amb interradii
(1.3-2.5 jmi).
Dimensions (30 specimens): Equatorial diameter
40 (49) 58 urn.
Holotype: Preparation D503/1, 91.8 13.1,
Y.3571; Plate 2, figure 8. Approximately triangular
amb, 51 /um in diameter, with virtually straight sides
PALYNOLOGY, VOLUME 6 — 1982
and rounded apices, where the exine is 3.2 ^m thick
compared to 1.4 ^m interradially; laesurae sinuous,
extending to equator, with narrow, elevated lips;
exine laevigate.
Type locality: Locality 1.
Derivation of name: After Dr. Jan Muller of the
Rijksherbarium, University of Leiden, Holland.
Comparison: There is superficial similarity be-
tween Matonisporites mulleri sp. nov. and certain
fossil spores that have been attributed to such genera
as Cyathidites Couper 1953, Deltoidospora Miner
1935, and Cyathea Smith (e.g. by Martin, 1973). The
present species differs, however, in having sinuous,
lipped laesurae and differentially thickened exine.
Affinity: Possibly with either of the fern families
Matoniaceae or Dicksoniaceae (see Couper, 1958;
Dettmann, 1963).
Occurrence: Localities 1,3, and 5.
Genus Gleicheniidites Ross emend. Skarby 1964
Gleicheniidites circinidites (Cookson) Dettmann 1963
Plate 3, figure 1
Remarks: A few specimens only were encountered
of this species, which is widely known from Mesozoic
and Cenozoic strata (Martin, 1973, p. 6.).
Affinity: The spores of the modern fern Glei-
chenia microphylla R.Br. (i.e., G. circinata Sw. in
Harris, 1955, p. 70, pi. 2, fig. 17) are closely similar.
Occurrence: Locality 1.
Genus Polypodiaceoisporites Potonie 1951
ex Potonie 1956
Polypodiaceoisporites retirugatus Muller 1968
Plate 2, figures 1-7
Description: Spores radial, trilete. Amb subtrian-
gular with convex to almost straight sides and
rounded apices. Laesurae extend to or almost to
inner margin of cingulum: accompanied by conspicu-
ous lips, individually 3-7 ftm wide. Cingulum
laevigate, 3-6 jon wide, width + uniform on given
 Neogene Palynomorphs from Papua New Guinea
specimen. Distal sculpture (i.e., of non-cingulate
exine) consisting mainly of irregularly developed,
smooth, rounded rugulae, 1.5-8 /xm wide, together
with minor interspersed verrucae. The sculptural
elements average 2 /*m in height and are normally
absent or much subdued proximally.
Dimensions (15 specimens): Equatorial diameter
47 (56) 65 urn.
Remarks: Polypodiaceoisporites tumulatus Part-
ridge (in Stover and Partridge, 1973), from the late
Miocene of the Gippsland Basin, southeastern
Australia, is smaller (32-44 fim in diameter) with
slightly concave sides (equatorial outline) but is
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otherwise closely similar to Muller's species. Some of
the present specimens resemble Khan's (1976a)
species Cingulatisporites papuanus and C. ivirensis,
from the Upper Tertiary of Papua New Guinea, but
differ mainly with respect to sculptural details and
cingulum widths as specified by Khan.
Muller (1968) recorded P. retirugatus from the
Late Cretaceous to Early Tertiary of Sarawak.
Affinity: Muller (op. cit.) alluded to probable
affinities with the fern genus Pteris L., "the mislead-
ing (form-) generic designation notwithstanding"
(see also Kremp and Kawasaki, 1972, p. 42).
Occurrence: Localities 1,3, and 5.
Monolete Spores
Genus Laevigatosporites Ibrahim 1933
Laevigatosporites major (Cookson) Krutzsch 1959
Plate 3, figures 2, 3.
Remarks: As noted by Hekel (1972, p. 5), this
simple form category is common in the Tertiary, and
indeed is represented from the Carboniferous on-
wards.
Occurrence: Localities 1 and 5.
Genus Microfoveolatosporis Krutzsch 1959
Microfoveolatosporis sp.
Plate 3, figures 4-11
Remarks: Included here are small numbers of
39
monolete, sculptured spores with simple laesura and
variably thin- to thick-walled exine that is punctate to
negatively reticulate.
Affinity: Spores of this morphological type are
known to be produced by certain pterophytes of the
Family Schizaeaceae (Bolkhovitina, 1961; Dettmann,
1963, p. 87) and psilophytes of the Family
Psilotaceae (Huang, 1981).
Occurrence: Localities 1,3, and 5.
Genus Scolocyamus gen. nov.
Type-species: Scolocyamus magnus sp. nov.; here
designated.
Diagnosis: Spores bilaterally symmetrical, mono-
lete, acavate. Amb roundly elliptical to oval. Distal
and proximo-equatorial regions of exine sculptured
with prominent, discrete, elongate projections of
spine-like form having simple or, more typically,
branched apices.
Name derivation: Gr., skolos, thorn; Gr., kyamos,
bean. Gender masculine.
Comparison: Tuberculatosporites Imgrund 1960
is an acavate, apiculate, monolete spore genus, but
is characterized by a comprehensively developed
conate to spinose sculpture, the elements of which
are of much lesser magnitude (<3 /xm long on the
type-species, T. anicystoides Imgrund 1960). Echina-
tosporites Akyol 1964 differs from Scolocyamus gen.
nov. in having simple (unbranched) spines, typically
with distinctly expanded (mammoid) bases. Another
monolete apiculate genus, Echinoidites Gonzalez
Guzman 1967, has coarse, invariably simple spines of
regular form and disposition and is thereby distin-
guishable from Scolocyamus.
Affinity: As discussed below, the type-species
(Scolocyamus magnus sp. nov.) appears to be closely
comparable, if not identical, with the spores of the
present-day blechnaceaeous fern Stenochlaena areo-
laris (Harris) Copeland.
Scolocyamus magnus sp. nov.
Plate 4, figures 4-7; Text-figure 2
1975 [Dispersed spores attributed to] Stenochlaena
areolaris; Anderson and Muller, pp. 295-296,
pl. 1, fig. 1.
 40
Diagnosis: Spores bilateral, monolete. Amb ellip-
tical. In equatorial view, shape is piano- to (slightly)
concavo-convex (distal surface strongly convex).
Laesura accompanied by lip-like exinal folds; length
ca. one-half to three-quarters of spore length. Exine
thin (1.2-1.9 /xm), bearing conspicuous, elongate,
discrete sculptural projections, grossly spine-like in
form, well-spaced on distal and proximo-equatorial
regions and absent from bulk of proximal surface.
Projections circular in basal outline (diameter
2.5-7.5 /xm), tapering gradually to apices that are
simple (blunted-acute to slightly bulbous) or, more
usually, are irregularly branched (bifurcant to multi-
furcant), the ultimate tips being blunted. Spacing of
projections varies from 3.5 /xm to 16 /xm. Length of
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projections 5-17 /xm, the shorter ones generally sited
proximo-equatorially. Exine otherwise laevigate to
scabrate.
Dimensions (18 specimens): Equatorial diameter
(length) 78 (100) 126 /xm. Polar diameter 45 (57)
73 /xm.
Holotype: Preparation D502/2, 116.1 8.7,
Y.3595; Plate 4, figure 4. Shape nearly plano-convex
in equatorial aspect with distinctly convex distal
surface and slightly concave proximal surface; length
of spore 95 /xm, polar diameter 57 /xm; laesura 58 /xm
long, accompanied by exinal folds; exine 1.2 /xm
thick with elongate, stout, spine-like sculptural
elements projecting from distal surface and from
narrow proximo-equatorial region bordering
laevigate contact areas; spines 2.5-7.5 /xm broad
basally, 6-17 /xm long, spaced 3.2-13 /xm apart.
Type locality: Locality 5.
Derivation of name: Latin, magmts, large.
Remarks: Due to the thinness of the exine,
PLATE 4
1-3 Polypodiidites sp., X500. 1, Lateral aspect; D504/2, 90.7
17.2, Y.3592. 2, Distal focus; D503/1, 117.2 5.9, Y.3593. 3,
Proximal focus; D502/3, 100.3 13.7, Y.3594.
4-7 Scolocyamus magnus gen. et sp. nov. 4, Holotype, lateral
aspect, X75O; D502/2, 116.1 8.7, Y.3595. 5, Lateral aspect,
X750; D502/3, 95.5 7.9, Y.3596. 6, Lateral aspect, X1000;
PALYNOLOGY, VOLUME 6 — 1982
specimens of Scolocyamus magnus gen. et sp. nov.
tend to be broken or have a distorted or corroded
appearance. Isolated scraps of exine bearing the
characteristic sculptural elements (Text-Figure 2) are
not uncommon (e.g. Plate 4, fig. 7).
Spores of this distinctive morphological type have
been reported from the Miocene and Pliocene of
northwest Borneo (Anderson and Muller, 1975;
Morley, 1978), but not previously as fossils from
Papua New Guinea.
Affinity: Holttum's (1932, p. 252, fig. 3) line-
drawing of a spore of Stenochlaena areolaris (Harris)
Copeland is sufficient to indicate a relationship of
the spores described above with the latter species.
This relationship is underpinned by Anderson and
MuIIer's (1975) observations, stressing the "unique"
spore morphology of S. areolaris.
Stenochlaena is a genus of the fern family Blech-
naceae. According to Anderson and Muller (op. cit.),
S. areolaris is at present "an epiphyte of Pandanus in
the Philippines and New Guinea at an altitudinal
range of 150-1200 m but has not been recorded from
Borneo."
Occurrence: Locality 5.
Genus Stenochlaenidites Khan 1976
Stenochlaenidites papuanus (Cookson) Khan 1976
Plate 4, figure 8; Plate 5, figures 1-3
1957 Schizaea papuana Cookson (partim), p. 44;
pi. 8, figs. 8, 9, 11. (non pi. 8, figs. 10, 12)
1976 Stenochlaenidites papuanus (Cookson) Khan,
p. 759, fig. 11 [1976a].
Remarks: The exine of these monolete spores is
distinctively sculptured, outside contact areas, by
D502/S1/8, 109.7 16.4, Y.3597. 7, Detail of sculptural pro-
jections based on fragmented portion of distal surface,
X2000; D502/S1/11, 110.9 15.2, Y.3598.
Stenochlaenidites papuanus (Cookson) Khan 1976. Lateral
aspect, X1500; D502/S1/16, 108.9 16.2, Y.3599.
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Neogene Palynomorphs from Papua New Guinea

PLATE 4
 42
20-i
15-
10-
5-
Text-Figure 2. Sculptural elements of Scolocyamus magnus gen. et sp. nov.; lateral views.
verrucae that are mostly arranged uniserially to form
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crenate or knobbly ridges. In lateral aspect, the
spores are piano- to concavo-convex, 49-73 fira. long
(12 specimens measured).
Khan (op. cit.) recorded S. papuanus from the late
Miocene to Holocene of the Central Delta region of
Papua New Guinea. Morley (1978, fig. 2, with 1980
hand-written amendment) noted its presence in late
Miocene and Pliocene sediments of Borneo.
Affinity: This form-species appears to be identical
with spores produced by Stenochlaena laurifolia
Presl (see Holttum, 1932, p. 252, fig. 2; Morley,
1978), a fern of the family Blechnaceae. S. laurifolia
is extant in the Philippines and Papua New Guinea
but not in Borneo (see Morley, op. cit, fig. 9). The
African species, S. tenuifolia (Desv.) Moore also
produces spores of similar morphology (Erdtman,
1957, p. 94, fig. 183; Erdtman and Sorsa, 1971, p.
175).
Occurrence: Locality 5.
Genus Polypodiidites Ross ex Couper, 1953
Polypodiidites usmensis (van der Hammen)
Hekel 1972
Plate 5, figures 4-7
Remarks: The verrucae of these monolete spores
are mainly discrete, often cone-like, and tend to be
randomly dispersed on the distal and proximo-
equatorial regions. Length of specimens varies from
43 to 97 iim, mean 66 jtm (20 specimens measured);
shape piano- to slightly concavo-convex. For
synonymy listing see Hekel (1972, p. 6), who also
noted possible conspecificity with Polypodiidites per-
verrucatus Couper 1953.
PALYNOLOGY, VOLUME 6 — 1982
P. usmensis has been recorded hitherto from the
Eocene to Holocene of northern South America,
Nigeria, and Borneo (Germeraad et al., 1968;
Anderson and Muller, 1975; Morley, 1978), Neogene
of Queensland (Hekel, 1972), Pliocene of Papua New
Guinea (Cookson, 1957), and Miocene of south India
(Rao and Ramanujam, 1978).
Affinity: The general category of verrucate or
gemmate, monolete spores is a common one among
the Pterophyta. However, as noted by Germeraad et
al. (1968, p. 292; pi. 2, fig. 4), this particular form, in
which the sculptural elements are mostly well-
separated from each other, is diagnostic for the
climbing swamp fern Stenochlaena palustris (Burm.
f.) Beddome of the family Blechnaceae (see also
Holttum, 1932, p. 252, fig. 1).
S. palustris is the most widespread of the three
extant Indo-Malesian species of Stenochlaena,
occurring in tropical Asia, Australia, and Polynesia
(Holttum, 1932).
Occurrence: Localities 1,3, and 5.
Polypodiidites sp.
Plate 3, figures 12, 13; Plate 4, figures 1-3
Description: Spores radial, trilete. Outline, in
polar view, broadly elliptical; in lateral view, plano-
convex. Laesura simple or occasionally with very
narrow lips, straight, length about 0.5-0.6 of spore
length. Verrucate sculpture developed equatorially
and distally, consisting of close-spaced, smooth,
rounded verrucae that are usually separated by a fine
negative reticulum; verrucae irregularly rounded to
polygonal in basal outline (diameter 1.5-18 pm),
0.5-3 pm high. On proximal surface, contact areas
are usually devoid of sculpture or bear only minor,
 Neogene Palynomorphs from Papua New Guinea
poorly defined elevations. A few specimens, how-
ever, display comprehensively verrucate exines.
Exine, excluding sculptural projections, 1.9-3.8 ftm;
normally thicker disto-equatorially than in contact
areas.
Dimensions (40 specimens): Equatorial diameter
— length 50 (65) 94 /xm, breadth 36 (45) 68 /xm.
Remarks: These spores could not be assigned un-
equivocally to any one previously described form
species because of the continuous nature of the mor-
phological variation among them. It can be noted,
however, that there are included here specimens com-
parable to such species as: Polypodiidites inanga-
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huensis Couper 1953 (p. 29; pi. 2, fig. 16), Verrucato-
sporites speciosus Harris 1965 (pp. 83-84; pi. 24,
figs. 8-10), and Polypodiidites sp. of Martin 1973 (p.
14; figs. 53, 54).
Affinity: Botanical relationships appear to be with
the polypodiaceaeous ferns, such as Microsorium
diversifolium (Willd.) Copeland (see Couper, 1960,
p. 39; pi. 1, figs. 5, 6) and Polypodium vulgare L.
(see Kremp and Kawasaki, 1972, p. 110).
Occurrence: Localities 1,3, and 5.
Bisaccate Pollen
Genus Lygistepollenites Stover & Evans 1973
Lygistepollenites florinii (Cookson & Pike)
Stover & Evans 1973
Plate 5, figure 8
Remarks: Several examples of L. florinii were
encountered (overall length 56-66 ftm). The species is
well-known from Tertiary sediments in the Australia-
New Guinea region (Cookson and Pike, 1953; Stover
and Partridge, 1973; Khan, 1976b).
Affinity: Cookson and Pike (1953) demonstrated
the natural affinity of L. florinii with Florin's (1940)
'Group B' category of the podocarpaceaean genus
Dacrydium Soland.
Occurrence: Locality 3.
Monocolpate Pollen
43
Genus Couperipollis Venkatachala & Kar 1969
Couperipollis spp.
Plate 5, figures 9-15; Plate 7, figure 13
Remarks: Included in this probable multispecific
category is a range of morphological types of mono-
sulcate pollen grains, elliptical in polar view
(maximum length 50-64 fim), with finely punctate-
reticulate, semitectate exines bearing ornament of
spinae to coni of sparse to fairly dense distribution.
Affinity: These forms appear to be allied to the
monocot family Liliaceae (cf. Astelia sp. and A.
cunninghamii Hook. f. in Couper, 1953, 1960).
Occurrence: Localities 1 and 3.
Dicolpate Pollen
Genus Dicolpopollis Pflanzl 1956
Dicolpopollis malesianus Muller 1968
Plate 6, figure 4; Plate 7, figure 4
Remarks: The present specimens conform with
Muller's (1968, p. 13; pi. 3, fig. 5) "very variable
species", although their size range (30 (37) 45 /xm; 30
specimens) diverges somewhat from that quoted by
Muller (22 (29) 35 (xm; 9 specimens). As stated by
Muller, Dicolpopollis maselianus "is known to occur
throughout the Tertiary up to the Recent."
Affinity: According to Muller (1968, p. 13)
affinity with the genus Calamus, of the palm sub-
family Lepidocaryoideae, is probable. As noted by
Sowunmi (1972, p. 28), the Calamus (palynological)
subgroup, which includes species of several lepido-
caryoid genera, differs from all other palms in pro-
ducing disulcate pollen grains.
Occurrence: Localities 1,3, and 5.
Tricolpate Pollen
Genus Perfotricolpites Gonzalez Guzman 1967
Type-species: Perfotricolpites digitatus Gonzalez
Guzman 1967; by original designation.
Perfotricolpites maculosus sp. nov.
Plate 6, figures 1-3
 44
Diagnosis: Pollen grains radiosymmetric, iso-
polar, prolate to subspherical, amb circular,
tricolpate. Colpi straight to slightly undulating,
reaching to within ca. 5-15 jon from poles. Exine
thick (4.2-6.5 ^im): nexine 0.9-1.7 ^m, sexine 3.3-4.8
£im. Columellae 1-1.4 ^m in diameter, 0.5-1 jtm
apart at their bases; abruptly and uniformly digitate
apically (branches <0.5 ^m wide and apart). Ratio
of simple to branched portions of columellae is
1.7-3.1. Surface of exine finely and uniformly
punctate.
Dimensions: Polar diameter 74-96 pm (10
specimens); equatorial diameter 58-78 ^m (3
specimens).
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Holotype: Preparation D504/2, 124.4 15.9,
Y.3616; Plate 6, figure 2. Prolate-subspherical, polar
diameter 94 pm, colpi 67 ^m long; exine 5-8 /xm
thick; sexine 4.8 ^m thick, consisting of columellae
1.3 ^m broad and 0.8 fim apart basally, apically
digitate to produce a very finely and densely
columellate, outer sexine layer 1.2 pm thick; surface
of exine finely and densely punctate.
Type locality: Locality 3.
Derivation of name: Latin, maculosus, dappled.
Comparison: Perfotricolpites digitatus Gonzalez
Guzman 1967 (see Germeraad et al, 1968, pp. 320,
322; pi. 12, fig. 10) is very similar to P. maculosus sp.
nov. but differs in being smaller and in having a
thinner exine.
Affinity: Clear resemblance exists with pollen
produced by certain species of certain genera
belonging to the eurypalynous family Convol-
PLATE 5
i-3 Stenochtaenidites papuanus (Cookson) Khan 1976; lateral
views, X500. 1, D502/3, 120.1 14.0, Y.3600. 2, D502/3,
108.6 16.3, Y.36O1. 3, D502/3, 93.7 16.7, Y.3602.
4-7 Polypodiidites usmensis (van der Hammen) Hekel 1972;
lateral views. 4, X500; D504/10, 107.9 17.9, Y.3603. 5,
X500; D503/1, 89.8 10.5, Y.3604. 6, X500; D5O3/1, 122.0
19.1, Y.3605. 7, X1500; D504/S1/10, 113.8 12.2, Y.3606.
8 Lygistepollenites florinii (Cookson & Pike) Stover &
Evans 1973. Distal focus, X750; D504/1, 94.1 18.4,
PALYNOLOGY, VOLUME 6 — 1982
vulaceae: see Germeraad et al. (1968, pp. 320-322),
Sengupta (1972, pp. 164-176), and Huang (1972, pp.
97-98).
Occurrence: Locality 3.
Genus Rostriapollenites
Venkatachala & Kar 1968
Type-species: Rostriapollenites kutchensis Ven-
katachala & Kar 1968; by original designation.
Discussion: Jansonius and Hills (1976 et seq.,
cards 167, 1604, 2441) have suggested possible
synonymy between this genus and one or both of
Areolipollis and Marginipollis which were estab-
lished by Clarke and Frederiksen (1968). Plancho-
niidites Khan 1976a, although specified as being "tri-
colporoidate or tricolpate," is obviously very closely
allied.
Rostriapollenites robustus sp. nov.
Plate 6, figures 5-10
Diagnosis: Pollen grains prolate, 3-syncolpate;
amb circular. Colpi crassimarginate, the mesocolpial
margins each marked by a distinct groove. Colpi
margins distinctly and abruptly thickened at poles
(5-7 ^m) to form distinctively beak-like oroid projec-
tions; elsewhere up to 2.7 ^m thick. Mesocolpial
exine 1.2-2 ^im thick, nexine: sexine thickness ratio
being approximately 1:1-2. Exine surficially
laevigate to scabrate but with subtectal areoloidate
pattern (most coarsely developed towards aperture
margins), due to somewhat irregular development of
sexinal columellae, 0.5-3 jan broad.
Y.3607.
9-15 CouperipoUis spp. 9, Median focus, X750; D504/2, 100.1
15.3, Y.3608. 10, Distal focus, X750; D504/3, 102.1 14.6,
Y.3609. 11, Median focus, X750; D504/20, 108.5 16.6,
Y.3610. 12, Distal focus, X750; D504/3, 100.3 15.5,
Y.3611. 13, Distal focus, X750; D504/8, 106.5 9.6, Y.3612.
14, Median focus, X750; D504/22, 105.5 15.0, Y.3613. 15,
Distal surface, X1500; D504/S1/26, 114.1 10.9, Y.3614.
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Neogene Palynomorphs from Papua New Guinea

PLATE 5
 46
Dimensions (28 specimens): Polar diameter 51
(58) 66 ftm; equatorial diameter 32 (39) 46 ^m.
Holotype: Preparation D504/1, 95.0 14.5,
Y.3619; Plate 6, figure 5. Grain prolate, diameter 63
ftm x 38 fttn; groove bordering incrassate colpate
margin up to 1.3 y,m wide, usually <0.5 /xm; meso-
colpial exine 1.9 /xm thick (nexine 0.6 ixm, sexine 1.3
ixxri); cuspidate projections (associated with colpi at
poles) result from greatly thickened exine (nexine 2.7
ftm, sexine 3.7 ^m); areoloidate exine pattern with
"verrucae" 0.7-2.5 (ivtx in diameter (actually breadth
of columellae) separated by fine "negative reticu-
lum", 1.2 ixtn or less wide.
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Type locality: Locality 3.
Derivation of Name: Latin, robustus, strong.
Comparison: Rostriapollenites robustus sp. nov.
differs from R. kutchensis Venkatachala & Kar 1968
by having a more subdued areoloidate pattern
together with grooved margins to the colpi; from
Marginipollis concinnus Clarke & Frederiksen 1968
chiefly by being appreciably larger with thicker meso-
colpial exine; and from Planchoniidites areolatoideus
Kahn 1976a in its more distinctly prolate shape, lesser
equatorial diameter, and more conspicuous cuspidate
polar projections.
Affinity: The pollen grains show clear morpho-
logical similarities to grains produced by the genus
Barringtonia of the family Lecythidaceae (or, more
restrictedly, of the family Barringtoniaceae): see
Payens (1967), Venkatachala and Kar (1968), Clarke
and Frederiksen (1968), Huang (1972, p. 136; pi. 83),
and Muller (1972, 1973). They conform most
obviously with the 'Barringtonia asiatica Type' of
Barringtonia pollen, specified by Muller (1972), on
PLATE 6
1-3 Perfotricolpites maculosus sp. nov., X500. 1, Polar aspect;
D504/19, 108.0 14.5, Y.3615. 2, Holotype, lateral aspect;
D504/2, 124.4 15.9, Y.3616. 3, Lateral aspect; D504/9,
109.5 10.5, Y.3617.
4 Dicolpopollis malesianus Muller 1968, X1250. D504/2,
123.0 13.9, Y.3618.
5-10 Rostriapollenites robustus sp. nov. 5, Holotype, lateral
PALYNOLOGY, VOLUME 6 — 1982
account of the presence of a groove demarcating the
apertural areas from the mesocolpia.
Occurrence: Localities 3 and 5.
Tricolporate Pollen
Genus Margocolporites Ramanujam
ex S.K. Srivastava 1969
Type-species: Margocolporites tsukadai Ramanu-
jam 1966; by subsequent designation of Srivastava
(1969, p. 984).
Margocolporites tricuneatus sp. nov.
Plate 7, figures 5, 6
Diagnosis: Pollen grains radiosymmetric, iso-
polar, oblate; subcircular to inter-semiangular in
polar view; 3-zonimargocolporate. Margocolpus
prominent, ca. 20 pm broad at equator, gradually
tapering to a rounded apex some 3-4 jxva from pole
and thus not joining with two other margocolpi.
Each margocolpus bordered by laevigate costa,
3.5-4.5 fan wide and 1.2-2 \aa high. Exine 2.5-3.5
jon thick; nexine thickness one-quarter to one-third
that of sexine, which is relatively coarsely columellate
outside costate borders but very finely so within.
Surface of margocolpus minutely and densely
punctate-vermiculate; elsewhere, exine surface is
reticulate to perforate (muri 1 fim. or less wide;
lumina subcircular to narrowly elongate in outline,
ca. 0.3 jim wide).
Dimensions (10 specimens): Equatorial diameter
47 (54) 62 nm.
Holotype: Preparation D504/1, 89.0 11.9,
Y.3629; Plate 7, figure 5. Amb inter-semiangular, 61
aspect, X750; D504/1, 95.0 14.5, Y.3619. 6, Polar aspect,
X750; D504/2, 109.8 7.6, Y.3620. 7, Lateral aspect, X75O;
D504/17, 110.8 11.9, Y.3621. 8, Lateral aspect, X75O;
D504/2, 89.3 19.7, Y.3622. 9, Off-polar aspect, X175O;
D504/S1/29, 115.2 10.5, Y.3623. 10, Lateral aspect,
X175O; D504/S1/17, 113.2 11,3, Y.3624.
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Neogene Palynomorphs from Papua New Guinea

PLATE 6
 48
/an in diameter; margocolpi 19 ^m broad equatorial-
ly, tapering uniformly to acutely rounded apices sited
ca. 3.5 /tm from pole, apertures extending 14-15 /mi
from equator towards pole; costae (defining margo-
colpi) 4 ^m wide, 1.8 ixm high; exine 3.5 fivn in thick-
ness (including 0.9 ^in-thick nexine), distinctly
reticulate-semitectate in mesocolpia, minutely per-
forate-tectate within costae.
Type locality: Locality 3.
Derivation of name: Latin, tri-, three-; cuneus,
wedge.
Comparison: Margocolporites vanwijhei Ger-
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meraad, Hopping, &Muller 1968 differs fromM. tri-
cuneatus sp. nov. chiefly by having shorter colpi,
thinner exine, and baculate margocolpi which, more-
over, have acutely pointed poleward termini.
Affinity: Amongst the Leguminosae, some genera
of the family Caesalpiniaceae produce pollen that are
morphologically akin to the grains described above.
This applies particularly to the pollen of present-day
Caesalpinia crista L. (see Huang, 1972, p. 142; pi. 86,
figs. 1-19) and C. coriaria Wild, (see Germeraad et
al, 1968, p. 342; pi. 18, figs. 4, 5).
Occurrence: Locality 3.
Tetracolporate Pollen
Genus Tetracolporopollenites Pflug & Thomson
in Thomson & Pflug 1953
Tetracolporopollenites sp.
Plate 7, figures 1-3
PLATE 7
1-3 Tetracolporopollenites sp., X750. 1, Polar aspect; D504/2,
108.0 7.8, Y.3625. 2, Lateral aspect; D504/2, 106.8 15.0,
Y.3626. 3, Lateral aspect; D504/2, 122.3 16.1, Y.3627.
4 Dicolpopollis malesianus Muller 1968, X1000. D5O2/3,
107.2 16.4, Y.3628.
5,6 Margocolporites Iricunealus sp. nov.; polar aspects, X750.
5, Holotype; D504/1, 89.0 11.9, Y.3629. 6, D504/1, 116.2
19.7, Y.363O.
7,8 Triporopollenites sp.; polar aspects, X750. 7, D504/3,
PALYNOLOGY, VOLUME 6 — 1982
Remarks: These pollen grains are prolate to sub-
spherical, 44-48 ^m in polar diameter, 33-41 pm in
equatorial diameter; four compound apertures (tetra-
colporate), longicolpate (length of colpi 25-30 jon),
pores lalongate to subcircular (diameter 2.5-5 /im)
and rimmed by thickened exine (3-4.5 ,»m thick);
exine two-layered, tectate to semitectate, sexine and
nexine mostly subequal in thickness (totalling 0.8-1.5
^m away from apertures), surface scabrate-punctate.
The specimens described above were insufficient to
serve as a basis for a new species. The generic desig-
nation applied here follows Potonie (1960, p. 110)
who indicated that Tetracolporopollenites is a senior
synonym of Sapotaceoidaepollenites Potonie, Thom-
son, & Thiergart 1950 ex Potonie" 1960 (see also Jan-
sonius and Hills, 1976, card 2500).
Some resemblance exists between the present speci-
mens and Sapotaceoidaepollenites rotundus Harris
1972, but the former are generally larger with thinner
exine and smaller ora.
Affinity: The botanical affinity is clearly with the
stenopalynous family Sapotaceae; see Erdtman
(1952, p. 397), Huang (1972, p. 217; pi. 140),
Anderson and Muller (1975, p. 308; pi. 2, fig. 10),
and Martin (1978, p. 188; figs. 6C, D).
Occurrence: Locality 3.
Genus Jandufouria
Germeraad, Hopping, & Muller 1968
Cf. Jandufouria sp.
Plate 7, figures 9-11
Remarks: Four specimens only of these tetracol-
poroidate grains were encountered, 40-44 jun in
105.4 17.0, Y.3631. 8, D504/21, 104.2 14.6, Y.3632.
9-11 Cf. Jandufouria sp. 9, Polar aspect, X75O; D504/2, 106.9
17.5, Y.3633. 10, Lateral aspect, X750; D504/23, 107.9
12.2, Y.3634. 11, Polar aspect, X2000; D504/S1/16, 112.9
11.2, Y.3635.
12 Epiphyllous fungal 'germling', X1000. D502/2, 125.9 17.4,
Y.3636.
13 Couperipollis sp.; disto-equatorial aspect, X1500.
D504/S1/11, 114.1 12.3, Y.3637.
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Neogene Palynomorphs from Papua New Guinea

PLATE 7
 50
equatorial diameter, and with subcircular amb. Pores
not clearly delimited, lolongate; colpilong, extending
two-thirds to four-fifths of distance from equator to
poles. Exine tectate-minutely perforate, 2.7-4.5 (xm
thick; ratio of nexine to sexine thicknesses is 1-1.5:1.
Exine of mesocolpus gradually thinned to colpi
margins.
Affinity: Germeraad et ah, (1968, p. 343)
indicated probable affinity of their Jandufouria
seamrogiformis (type-species) with the genus Cato-
stemma (family Bombacaceae); but this does not
necessarily apply to the present specimens that are
not indubitably assignable to Jandufouria.
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Occurrence: Locality 3.
Triporate Pollen
Genus Triporopollenites Pflug & Thomson
in Thomson & Pflug 1953
Triporopollenites sp.
Plate 7, figures 7, 8
Remarks: The few, relatively large triporate grains
included here exhibit the following characters. Amb
circular-subcircular; pores slightly aspidate, 3.8-6
pirn in diameter, rims thickened (3.5-4.5 \xvd); exine
elsewhere 1.8-3.2 pun thick, tectate; sexine and
nexine of approximately equal thickness (except
about pores where nexine is much thicker); sexine
surficially laevigate to scabrate; equatorial diameter
48-56 nm (5 specimens).
Affinity: Botanical relationships are unclear, but
somewhat similar pollen is produced by the
Ulmaceae.
Occurrence: Locality 3.
Discoidal invaginated microfossils
Epiphyllous fungal "germlings"
Plate 7, figure 12
Remarks: Two specimens only of these micro-
fossils, 25 ftxa. and 27 pim in diameter, were
encountered. They exhibit a circular outline with
regular "short-long" invaginations around the
PALYNOLOGY, VOLUME 6—1982
margin, and are clearly identifiable as "germlings"
of epiphyllous fungi. In detail, the specimens
conform with Lange's (1976) Grade VI or Grade V
(sensu lato) of "germlings" said to be characteristic
of rainforest vegetation (Type F of Lange, 1976, pp.
160-161).
Affinity: Fungal family Microthyriaceae.
Occurrence: Locality 5.
DISCUSSION
Table 1 presents a summary of the occurrence of
the palynomorphs variously identified in the three
productive carbonacous samples, together with
botanical affiliations, where possible to the level of
family. In some instances, more precise affinities are
evident (see systematic section).
In all of the samples, the lower vascular plants are
overwhelmingly represented both qualitatively and
quantitatively. Of these, the pterophytes (ferns) were
easily the most prominent palynological contrib-
utors. Thus it appears that the peat swamp vegetation
was dominated by the ferns, among which the
families Polypodiaceae, Schizaeaceae, Matoniaceae
(or Dicksoniaceae), and Blechnaceae were well-
represented. The last-named family is evidenced
palynologically by three miospore species that are
confidently assignable, respectively, to all three
extant Indo-Malesian species of the genus
Stenochlaena: S. palustris (the climbing swamp fern;
from all three localities), S. laurifolia (locality 5), and
S. areolaris (locality 5). However, the majority of the
fern spores present are not, from current evidence,
attributable to living species or genera, nor in some
cases unequivocally even to particular families. The
lycopods were evidently only minor components of
the peat-forming vegetation.
Palm pollen of the distinctive Calamus type is con-
sistently represented in the studied assemblages. As
noted by Haseldonckx (1977, p. 234), Calamus
species today inhabit swamp perimeters beyond
saline influence. Occurring prominently at two
localities (3 and 5) are pollen derived from the genus
Barringtonia, at present a shrub or small tree in
tropical, freshwater swamps of Africa, southeast
Asia, northern Australia, and the Pacific islands
 Neogene Palynomorphs from Papua New Guinea 51

TABLE 1

SUMMARY OF OCCURRENCES AND BOTANICAL AFFINITIES OF PALYNOMORPH TAXA

RECORDED HEREIN
LOCALITY BOTANICAL AFFILIATION
FOSSIL SPECIES
1 3 5 Oivision Family
Biretisporites huonensis a a a Pterophyta

Baculatisporites scabridus a Bryophyta or Pterophyta

Converrucosisporites ponderosus a Pterophyta

Foveosporites sp. A © a Lycopodophyta Lycopodiaceae

Foveosporites sp. B a Lycopodophyta or Pterophyta

Rugulatisporites trophus 9 Lycopodophyta Lycopodiaceae

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Cf. Rugulatisporites sp. a Pterophyta (?)

Matonispori.t&s mulleri Pterophyta

Gleicheniidites circinidites e Pterophyta Gleicheniaceae

Polypodiaceoisparites retiruyatus 0 a a Pterophyta Pteridaceae

Laevigatosporites major a 9 Arthrophyta or Pterophyta

Microfoveolatosporis spp. & e 9 Pterophyta Schizaeaceae

Scolocyamus magnus a Pterophyta Blechnaceae

Stenochlaenidites papuanus a Pterophyta Blechnaceae
Poly pod1idltes usmensis 9 o a Pterophyta Blechnaceae
Polypodiidites sp. e a a Pterophyta Polypodiaceae
Lygistepollenites florxnii a a Coniferophyta Podocarpaceae

Couperipoll is spp. © e Anthophyta Liliaceae

Dicolpopollis malesianus a a 9 Anthophyta Arecaceae
Perfotricol pites maculosus a Anthophyta

Rostriapollenites robustus a a Anthophyta Barringtoniaceae

Margocolporites tricuneatus a Anthophyta Caesalpiniaceae

Tetracolporopollenites sp. a Ar *"hophy ta Sapotaceae
Cf. Jandufouria sp. a Anthophyta Bombaceae (?)

Triporopollenites sp. a Anthophyta Ulmaceae (?)

Epiphyllous fungal 'germlings' a Ascomycota Microthyriaceae

(Payens, 1967). Other angiospermous groups
represented, albeit spasmodically, are the families
Liliaceae, Convolvulaceae, Caesalpiniaceae, Sapot-
aceae, and possibly also Bombacaceae and
Ulmaceae.
No pollen grains of Nothofagus were encountered,
rather surprisingly in view of their reported
abundance (as the brassii type) in the Neogene of
Papua New Guinea (Khan, 1974). Other notable
absentees from the samples studied include the man-
groves, the Sonneratiaceae, Myrtaceae, and Gunner-
aceae (cf. Cookson and Pike, 1954; Khan, 1974). The
only indications of gymnosperms are a few bisaccate
grains of Dacrydium (Group B) recorded from
localities 3 and 5.
The palynological assemblages afford no positive
evidence of marine influence; i.e., in the form of
organic-walled marine phytoplankton or pollen from
known halophytes. In fact, fragments of the fresh-
water alga Pediastrum occur, though infrequently, in
the studied samples. This is perhaps surprising in
view of the dominantly marine facies in the Pindiu
area and the foraminiferal content of samples from
localities 1 and 3.
 52
The available evidence suggests that the spore-
pollen assemblages have derived autochthonously
from a tropical, freshwater, peat swamp vegetation
of relatively low diversity. If so, the pterophytic
component of the vegetation would have effectively
masked, or at least rendered inconspicuous, palyno-
logical contributions from most other taxonomic
groups.
Especially noteworthy is the virtual absence of
wind-transported pollen including elements of the
Fagaceae {Nothofagus, as mentioned above) and
Podocarpaceae which have formed rainforest
canopies from at least the late Miocene onwards in
the New Guinea region (Khan, 1974). It follows,
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then, that the studied palynofloras reflect a special-
ized ecosystem, hardly representative of the overall
contemporary vegetation of the region.
The generally similar complexion of the palynolog-
ical assemblages from the three samples implies that
there may be no significant age disparity between
them. Consistently, and often abundantly repre-
sented are the following form-species: Biretisporites
huonensis, Matonisporites mulleri, Polypodiaceoi-
sporites retirugatus, Microfoveolatosporis spp.,
Polypodiidites usmensis, Polypodiidites sp., and
Dicolpopollis malesianus. Three other forms are
represented in two samples; viz., Foveosporites sp.
A, Couperipollis spp., and Rostriapollenites
robustus. With a few exceptions, the main composi-
tional differences among the samples are in the inci-
dence of the rarer forms, such as Rugulatisporites
trophus, Baculatisporites scabridus, Converrucosi-
sporites ponderosus, Gleicheniidites circinidites,
Lygistepollenites florinii, Cf. Jandufouria sp., and
Triporopollenites sp.
The palynoflora affords no really precise,
independent testimony as to the age or ages of the
samples, apart from being generally indicative of a
Neogene (Miocene or Pliocene) age, in accord with
the established gross age of the sediments exposed in
the Pindiu area. This indication is based mainly on
the fern spore species Scolocyamus magnus and
PALYNOLOGY, VOLUME 6 — 1982
Stenochlaenidites papuanus, both of which are
recorded from locality 5. Scolocyamus magnus has
previously been reported from the Miocene-Pliocene
of northwest Borneo; and its parent fern species,
Stenochlaena areolaris, is extant in the Philippines
and Papua New Guinea. So, too, is Stenochlaena
laurifolia, the miospore of which (Stenochlaenidites
papuanus) has been reported hitherto from the late
Miocene to Pliocene of northwest Borneo and from
the late Miocene to Holocene of Papua New Guinea.
The other Stenochlaena derivative (namely of S.
palustris), Polypodiidites usmensis, is present in all
three samples, and is known to be much longer
ranging, dating back to Eocene (in northwest
Borneo). The other miospore taxa of the assemblages
are mainly either new, and therefore of unknown
stratigraphic distribution, or have known ranges
extending back at least as far as the Paleogene.
ACKNOWLEDGMENTS
Dr. David Haig of the University of Papua New
Guinea is thanked for provision of the samples that
form the basis of this study. The research was
considerably aided by helpful comments and
suggestions from the following: Dr. Jan Muller
(Rijksherbarium, Leiden), Alan D. Partridge (Esso
Australia Limited, Sydney), Dr. Helene Martin (Uni-
versity of New South Wales, Sydney), Dr. David
Pocknall (New Zealand Geological Survey, Lower
Hutt), Peter Price (AAR Limited, Brisbane), and Dr.
Trevor Clifford (Department of Botany, University
of Queensland). My wife, Dr. Mary Dettmann,
offered much useful advice during the investigation.
Mr. J.V. Hardy of the Electron Microscope
Centre, University of Queensland, expertly aided the
scanning electron microscopy. In the Department of
Geology and Mineralogy, Mrs. D. Phipps was
responsible for laboratory processing of samples; Dr.
G. Khorasani provided coal-petrographic informa-
tion; Mrs. E. Burdin drafted the text-figures; and
Miss R. Austin typed the manuscript.
 Neogene Palynomorphs from Papua New Guinea
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