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A review of the use of non-pollen palynomorphs in
palaeoecology with examples from Australia
Ellyn J. Cook
, Bas van Geel
, Sander van der Kaars
& Jan van Arkel
School of Geography and Environmental Science, PO Box 11A, Monash University, Victoria,
3800, Australia
Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Science Park
904, 1098 XH Amsterdam, The Netherlands
Available online: 10 Jun 2011
To cite this article: Ellyn J. Cook, Bas van Geel, Sander van der Kaars & Jan van Arkel (2011): A review of the use of non-
pollen palynomorphs in palaeoecology with examples from Australia, Palynology, 35:2, 155-178
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A review of the use of non-pollen palynomorphs in palaeoecology with examples from Australia
Ellyn J. Cook
*, Bas van Geel
, Sander van der Kaars
and Jan van Arkel
School of Geography and Environmental Science, PO Box 11A, Monash University, Victoria 3800, Australia;
Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam, Science Park 904, 1098 XH Amsterdam,
The Netherlands
Records of the past climate and vegetation of Australia are frequently constructed using data generated from the
analysis of pollen and pteridophyte spores alone, or in association with sedimentology. We demonstrate that
the organic residue prepared for pollen analysis yields other organic-walled microfossils that can be used to provide
additional and independent palaeoenvironmental information. These non-pollen palynomorphs (NPP) include
microscopic remains of algae, cyanobacteria, fungi, insects, other invertebrates and cormophytes. Our study of the
NPP from two Late Quaternary lake records from western Victoria, Australia, provided additional information on
water quality, salinity, depth, temperature and nutrient levels to the general environmental interpretation derived
from pollen data. From a review of the ecological preferences of taxa, and comparison of the NPP results with pollen
and spore curves from the lake records, ecological indicator values were derived. The study confirms the utility of
NPP in enhancing environmental reconstructions in Australia, and encourages their routine examination in
palynological studies.
Keywords: non-pollen palynomorphs; Quaternary; Australia; indicator values
1. Introduction
1.1. Background
Over the past three de cades a sustained effort has
been made by many workers to describe and, where
possible, identify non-pollen microfossils encountered
in palynological studies. Most work has been
undertaken in northwest Europe, where van Geel
(e.g. 1972, 1986, 2001) and others (e.g. Bakker and
van Smeerdijk 1982; van der Wiel 1982; Kuhry 1985;
Haas 1996; Guy-Ohlson 1998; Lo
ez et al.
1998) have systematically developed the practice of
examining all fossils from a wide variety of sediment
types. The increased focus has brought a greater
appreciation of the indicator role that non-pollen
palynomorphs (NPP) can play in corroborating or
refining ecological interpretations. In early work in
the Netherlands, Pals et al. (1980) used pollen
together with remains of algae, fungi and plant
macrofossils to provide indications of the degree of
salinity, alkalinity and eutrophication of Klokkeweel
Bog, a site nearby the Bronze Age settlement of
Hoogkarspel-Watertoren (c. 1250–900 BC). Changes
in the fossil assemblages recorded the extent to
which humans altered the vegetation, landscape and
water quality at the site and provided evidence for
the abandonment of all settlements in the region
by *600 BC.
In the Sonoran desert in the USA and Mexico,
Davis et al. (2002) recorded elevated levels of spores of
the dung fungus Sporormiella at the introduction of
domesticated grazing animals during the historical
period. In the same study, increases in spores from the
fungal saprophyte Tetraploa indicated an increased
accumulation of plant debris which, together with
decreased charcoal levels and an increase in the
presence of woody taxa, identified the cessation of
seasonal burning practices by native Indians at contact
with European settlers. The reduction in fire frequency,
and increased amount of decaying vegetation, ex-
plained how the transformation of wetland sediments
from silt to peat developed. Previously, seasonal
burning had removed senescent plant material which,
through ash, had returned nutrients to the system.
More, recently, detailed investigations demonstrat-
ing the value of NPP have extended to Madagascar
(Burney et al. 2003), Mexico and the USA (Almeida-
Lenero et al. 2005; Gill et al. 2009), Brazil (Medeanic
2006) and India (Limaye et al. 2007). Most recently,
studies have been presented from Venezu ela (Montoya
et al. 2010), Canada (Graf and Chmura 2006;
McAndrews and Turton 2010), central and southern
*Corresponding author. Email:
Vol. 35, No. 2, December 2011, 155–178
ISSN 0191-6122 print/ISSN 1558-9188 online
Ó 2011 AASP The Palynological Society
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Figure 1. The location of lakes Bolac and Turangmoroke in
western Victoria, Australia.
Europe (Cugny et al. 2010; Mudie et al. 2010), and
Greenland (Gauthier et al. 2010). These studies
continue to make it clear that NPP should be routinely
analysed as palaeoecological proxies. This is especially
so given that many NPP come from algal and fungal
taxa more local in source and distribution than most
pollen, and that may be present when pollen produc-
tion, deposition and/or preservation are limited or
1.2. Rationale
The present study arose in response to a significant
presence of a wide diversity of non-pollen objects in
microscope slides prepared for routine pollen analysis
of material from lunette-lake systems Bolac and
Turangmoroke within the drier part of western
Victoria, Australia (Figure 1). The complete palaeoe-
cological records are present ed in Cook (2009) with the
aim of this paper being to specifically report on the
results of the first comprehensive examination of NPP
from Australia.
2. Regional setting
Lake Bolac (37842
S, 1428 45
E, *90 km inland,
220 m above sea level) covers *1460 ha and is one of
the largest lakes on the Western Plains. A study of the
lake chemistry by Khan (2003) showed the mean secchi
depth is 25.4 cm (low light penetration), salinity
4.0% (brackish) and pH 8.6 (alkaline). The lake is
supplied by Fiery Creek with a total catchment area
of *1000 km
, and drained at the south by Salt Creek
which in turn flows into the Hopkins River which itself
flows into the sea just southeast of Warrnambool. A
dam on Salt Creek raised around 1975, makes the
present average (artificial) water depth of Lake
Bolac *1.5 m (Banfield and McKenzie 1995). Lake
Turangmoroke lies 2 km to the east of Lake Bolac,
immediately north of Fiery Creek, and covers 93 ha.
The lake is supplied by a small catchment of a few
hundred hectares limited to the basalt plain to the
north of the lake and its present average water depth is
1m. Lake Turangmoroke is saline–hypersaline
(48.5%: Land Conservation Council Victoria 1980).
The two lakes are not naturally connected to each
other. In the early 1900s, a channel dredged between
Lake Turangmoroke and Fiery Creek directed the
Turangmoroke outflow into Bolac, reducing the
salinity of Lake Turangmoroke but possibly increasing
the salinity of Lake Bolac (Crowley and Kershaw
1994). Oscillating lake levels during the last glacial–
interglacial transition resulted in the form ation of
lunettes (longitudinal dunes) that border each of these
lakes. The mean annual rainfall at the site is 540.2 mm
of which 103.20 mm falls in summer and 154.9 mm
falls in winter. Rainfall exceeds evaporation from April
to October (Land Conservation Council 1980). The
mean annual temperature at Hamilton, the closest
weather station in proximity and elevation (*79 km to
the west), is 13.38C. Temperatures vary from a mean
daily maximum of 25.78C in January and February to
128C in July and a mean daily minimum of 128Cin
February to 4.58C in July.
3. Materials and methods
3.1. Field techniques
The Lake Turangmoroke record is composed from two
cores. A 4.29 m core (LTC1) was recovered from the
near-centre of the lake using the percussion coring
method outlined in Cook (2009) and a 1.29 m core
(LTC2) taken near the western shore of the basin using
a PVC pipe-in-pipe technique after that of Tratt and
Burne (1980), to attain and integrate data on local
shore conditions into the longer record from the deeper
part of the lake. A 3.0 m core (LB) was recovered from
the near-centre of Lake Bolac by percussion coring
(Cook 2009).
156 E.J. Cook et al.
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3.2. Analytical methods
Samples were taken at 2–3 cm intervals from LTC1,
3 cm intervals from LTC2 and 3–5 cm intervals from
LB. Extraction of organic microfossils was as outlined
in Cook (2009). NPP and pollen were identified
at 6 1000 magnification and counted at 6 400 mag-
nification using a Zeiss Axiolab at Monash University
and a Zeiss Axiostar at the University of Amsterdam.
Most samples were counted to a minimum of 250
dryland pollen grains. Where pollen concentrations
were low or the pollen poorly preserved, either a
minimum of 100 dryland grains was counted or the
entire organic residue was analysed. Samples with
fewer than 50 dryland grains were restricted to the
basal parts of LTC1 and LB and are shown by sections
shaded with grey on the diagrams.
Usage of the term ‘Type’ follows van Geel (1978).
There is no taxonomic implication intended. Photo-
graphs of the NPP were taken using a Zeiss Photo-
microscope III fitted with Plan-NEOFLUAR 6 16,
6 40 and 6 63 variable immersion DIC lenses and a
Nikon D2Xs camera. In total, 38 palynomorphs
(‘Types’) are described, where possible illustrated and
ecological information and indicator values given. The
Types include 24 palynomorphs that were identified,
two as yet unidentified Types recorded as Type 180
(van Geel et al. 1983) and Type 209 (van Geel et al.
1989) and 12 newly recorded Types that remain
4. Presentation of palynomorphs recorded and
discussion of indicator values
4.1. Presentation of NPP results
The frequencies of occurrence of all NPP taxa recorded
in the cores are presented in Figure 2 (LTC1), Figure 3
(LTC2) and Figure 4 (LB). Percentages for all NPP
taxa are calculated relative to the dryland pollen sum
used in Cook (2009). Figures 2–4 were produced using
TILIA (version 2.0.B.4: Grimm 1990, 1991), and
TGView (version 2.0.2) using the zonation derived by
Cook (2009). NPP are arranged in the following
biological groups: algae, aquatic invertebrates, fungi
and other palynomorph Types. The following remain
ungrouped: Anthoceros type, Bryophyte type and
charcoal identified as charred Poaceae epidermal
material. Shading of some curves indicates an exag-
geration of 6 5 as a visual aid. Core lithology is shown
on each diagram. Summaries of major ecological
groups are shown to illustrate changes in vegetation.
The chronology for the records follows that establis hed
in Cook (2009), based on AMS radiocarbon analysis of
pollen concentrates and OSL analyses of quartz from
the 60–90 mm or 90–125 mm fractions.
4.2. Reconstructed environmental history from Lakes
Bolac and Turangmoroke
The palaeoenvironmental history of the drier part of
western Victoria through the past *90,000 years
discerned from the LTC1, LTC2 and LB records by
Cook (2009) is summarised here to provide a context in
which to place the NPP finds. The regional vegetation
during marine isotope stage (MIS) 5.1 was composed of
open woodland, dominated by Allocasuarina luehmannii
type with a diverse understorey. Rainfall is likely to have
been more equally distributed than at present given the
preference of Allocasuarina luehmannii for slight winter
to weak summer precipitation maxima today (Doran
and Hall 1983). During that time, Lake Turangmoroke
held freshwater of variable depth. A shift towards
reduced moisture availability indicated that at least
some part of MIS 4 is contained in the record but owing
to variable pollen representation and the frequency with
which the lake may have dried, it is difficult to determine
how much of this period is represented. Pollen rep-
resentation during mid MIS 3 suggests dryland vegeta-
tion similar to MIS 5.1 but more variable lake levels. A
change to open grassland-steppe occurred soon
after *47,000 years ago, and lake levels fluctuated
considerably before Lake Turangmoroke became shal-
low and saline. Throughout MIS 2 the dryland flora was
characterised by open grassland-steppe with almost no
trees while aquatic flora reflected further declining lake
levels and increasing salinity. Driest conditions, indi-
cated by deflation of lake sediments to the lunette, are
dated to between *18,000 and *11,000 cal yr BP. The
early Holocene was dominated by Allocasuarina verti-
cillata type until partial replacement by Eucalyptus
*8000–9000 cal yr BP when the pre-European vegeta-
tion cover was established.
4.3. Description, illustration and indicator value of
non-pollen palynomorphs identified
Details on life cycles, habits and environmental
conditions have been compiled here to mak e informa-
tion easily accessible, but direct use of the cited works
is recommended for greater depth, more detailed
morphological descriptions and additional illustra-
tions. Additional references for each palynomorph
type can be found in Cook (2006). Morphological
descriptions presented here incorporate observations
of fossil material examined in this study, and informa-
tion from published sources.
4.3.1. Chlorophyta (green algae)
The cell walls of the Chlorophyta are composed of
sporopollenin (Zetsche and Vicari 1931), the same
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158 E.J. Cook et al.
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material as that of pollen grains and fern spo res,
making them similarly resistant to destruction during
fossilisation and chemical laboratory processing.
Botryococcus (Plate 1, figure 1)
Description. Individual cells of length 5–15 mm and
width 2.5–6 mm, angular or spherical. Densely packed
colonies of 10–100 mm are held together by mucilage
slime with cells branching from centre.
Remarks. Colonies of the planktonic green alga genus
Botryococcus are of common occurrence as fossils in
material from the Late Precambrian to the present day
(e.g. Koma
rek and Marvan 1992; Guy-Ohlson 1992,
1998; Taylor and Taylor 1993; Sandiford et al. 2002).
Botryococcus is most often found in freshwater fens and
lakes as the cosmopolitan B. braunii (Batten and Grenfell
1996) but can also inhabit temporary ponds, pools,
Round 1965; Davis et al. 1977; Graham and Wilcox
coccus is much more adapta ble to brackish waters than
other Chlorophyceae algae (Blackburn and Temperley
1936). Moore and Carter (1923) recorded Botryococcus
in both fresh and saline waters (up to 9.4%)atGrand
Coulee in North Dakota and De Deckker (1988)
reported collection of B. braunii in southeastern Australia
from water bodies with salinities up to 20%.
Turbidity and nutrient status of the wat er also
appear to exert significant control over the prolifera-
tion of Botryococcus. In New South Wales, Dulhunty
(1944) found that Botryococcus preferred quiet, rela-
tively sediment free waters without subaerial plants but
in close enough proximity to shore for rotting
vegetation to be a supply of nitrogen for growth. In
studies of extant colonies in western Victoria, Yezdani
(1970) found that Botryococcus preferred oligotrophic
conditions but would tolerate eutrophic conditions.
Haas et al. (2007) found that increases in Botryococcus
resulted from eutrophication linked to human impact.
A preference for open water explains the inter-
pretation made in some palaeoecological records that
Botryococcus prefers deep water. Deeper water selects
against an excess of aquatic vegetation and suspended
sediment, ensuring availability of light and a relatively
low level of nutrients. At Tulare Lake in California,
Davis (1999) identified a rise in Botryoc occus
coincident with a decline in littoral vegetation as
indicative of open water and high lake levels. At Lake
George in New South Wales, Singh et al. (1981)
considered Botryococcus indicative of permanent open
water but noted that, on the basis of coexisting pollen
evidence, Botryococcus peaks occurred at variable
lake depths. Therefore, early observations by Black-
burn and Temperley (1936) that highest Botryococcus
occurrences were in samples with low amounts of other
freshwater algae, pollen and spores meant shallow
lakes, might be equally well explained by greater lake
depths resulting in less aquatic vegetation, particularly
in the littoral zone.
In the palaeoecological record LTC1, high frequen-
cies of Botryococcus,togetherwithCosmarium, Hydro-
charitaceae, My riophyllum sp., Myriop hyllum muelleri
and Pedia strum occurred during the periods identified as
Lake Turangmoroke was of va riable depth, clear and
received regu lar freshwater input (zones 5–8 , Figure 2).
A subsequent decline in Botryococcus an d corresponding
rise in saline tolerant Ruppia megacarpa type during MIS
2 (zone 4, Figure 2) reflected declining lake levels as well
as increasing salinity. Myriophyllum sp. and
Myriophy l-
lum muelleri can survive temporarily on wet mud left by
receding water levels (Aston 1973) and Botryococcus can
inhabit temporary ponds an d wet mud (Wes t and
Fritsch 1927; Graham and Wilcox 2000), thus their
reduction is not only a function of lowered water levels.
Continued repres entation after this time of Botryococcus
in the LB rec ord without Ruppia megacarpa type
indicated that, although Lake Bolac beca me increasingly
saline, lev els did not reach those o f Lake Turan gmoroke
that fluctuated from sa line to hypersal ine.
Desmidiaceae (Plate 1, figure 2)
Description. Cosmarium consists of two identical parts
(semi-cells). Cell length *15–160 mm and width 15–
65 mm. Wall smooth or ornamented with granules. The
centre of the cell is deeply incised where semi-cells join.
Remarks. Desmids are green algae which are basically
unicellular. Sexual reproduction is by conjugation
(Prescott 1948). Mainly found in freshwater environ-
ments, desmids are usually indicative of clean water as
under nutrient rich conditions they cannot grow as fast
as other algae (Blackmore 1984). They prefer low
Figure 2. (a) Palaeoecological diagram from Lake Turangmoroke core 1 (LTC1) showing changes in major dryland pollen
groups and aquatic pollen taxa. Exaggeration of selected curves is 6 5. Shaded bands indicate sections of core containing
samples with fewer than 50 dryland grains. (b) Palaeoecological diagram from LTC1 (continued) showing changes in algal and
aquatic invertebrates. Exaggeration of selected curves is 6 5. Shaded bands indicate sections of core containing samples with
fewer than 50 dryland grains. (c) Palaeoecological diagram from LTC1 (continued) showing changes in representation of fungi,
other palynomorph Types and charcoal. Exaggeration of selected curves is 6 5. Shaded bands indicate sections of core
containing samples with fewer than 50 dryland grains.
Palynology 159
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Figure 3. (a). Palaeoecological diagram from Lake Turangmoroke core 2 (LTC2) showing changes in representation of major dryland pollen groups, aquatic pollen, algae
and aquatic invertebrates. Exaggeration of selected curves is 6 5. Shaded band indicates section of core containing samples with fewer than 50 dryland grains. (b)
Palaeoecological diagram from Lake Turangmoroke core 2 (LTC2) showing changes in fungi, other palynomorph Types and charcoal representation. Shaded band indicates
section of core containing samples with fewer than 50 dryland grains.
160 E.J. Cook et al.
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Figure 4. (a). Palaeoecological diagram from Lake Bolac core (LB) showing changes in major dryland pollen groups, aquatic pollen, algae and aquatic invertebrates.
Exaggeration of selected curves is 6 6.5. Shaded band indicates section of core containing samples with fewer than 50 dryland grains. (b) Palaeoecological diagram from
Lake Bolac core (LB) showing changes in fungi, Palynomorph Types and charcoal representation.
Palynology 161
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conductivity and, therefore, only two species are found
in saline waters (M. Dingley, personal communication,
2009). The largest desmid genus is Cosmarium , the
species of which occur in freshwaters (Prescott 1964).
In Australia it is esti mated that there are *218 species
of Cosmarium (Entwistle et al. 1997). In this study we
identified Cosmarium distichum, characteristic of al-
most still freshwater which is (moderately) nutrient
poor. Although it usually lives in shallow waters,
Cosmarium distichum can live in deeper waters if
Plate 1. Non-pollen palynomorphs identified. Core and sample depths for specimens shown indicated in brackets: LTC1 refers
to Lake Turangmoroke core 1, LTC2 refers to Lake Turangmoroke core 2, LB refers to Lake Bolac. Scale bar represents 20 mm.
Figure 1. Botryococcus (LB 201–202 cm). Figure 2. Cosmarium distichum (LTC1 45–46 cm). Figure 3. Pediastrum (LB 241–
242 cm). Figure 4. Debarya glyptosperma (LTC1 49–50 cm). Figure 5. Spirogyra (a: LTC1 49–50 cm; b: LB 1–2 cm). Figure 6.
Zygnema type (LTC1 309–310 cm). Figure 7. Gloeotrichia type (a–c: LB 121–122 cm with akinate (resting spore) clearly visible in
example 7a and 7b). Figure 8. Bryophyte type (LTC1 117–118 cm). Figure 9. Anthoceros type (LTC1 133–134 cm). Figure 10.
Glomus (LTC1 45–46 cm). Figure 11. Neurospora crassa (LTC2 30–31 cm). Figure 12. Podospora type (LTC1 45–46 cm). Figure
13. Sporormiella type (a: LTC2 50–51 cm single cell; b: LTC1 9–10 cm fruiting body with cells). Figure 14. Tetraploa (LTC2 67–
68 cm).
162 E.J. Cook et al.
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floating fields of Selaginella are present (P.F.M.
Coesel, personal commun ication, 2009). In the lakes
studied here, the presence of Cosmarium distichum was
restricted to the LTC1 record in the periods identified
as MIS 5.1 (zone 7, Figure 2) and the European phase
(zone 1, Figure 2). It was generally associated with
floating aquatic Selaginella and with the resting eggs
of the planktonic rotifer Filinia, indicating relatively
deep freshwater in the older part of the record and
freshwater stream input in the European phase.
Pediastrum (Plate 1, figure 3)
Description. Radially symmetrical colonial green algae,
zoospores linked in platelike colonies (coenobia). Each
colony is *10–35 mm and may contain 4–128 cells.
Outer cells have 1–2 horns.
Remarks. Pediastrum colonies are commonly found
in freshwater phytoplankton communities (Sarmaja-
Korjonen et al. 2006). Worldwide the most common
and widely distributed species is P. boryanum which
occurs in a variety of waters (Jankovska
and Koma
2000; Koma
rek and Jankovska
, 2001). The salinity
tolerance in Australia determined from fossil repre-
sentation is *1.7% (Churchill et al. 1978) but from
study of extant phytoplankton communities in western
Victoria is *3.5% (Yezdani 1970).
Woolfenden (1993, 19 95) recorded Pediastrum bor-
yanum as characteristic of the littoral zone of oligo -
trophic waters. Fossil representations of Pedias trum
have also been correlated with deep, oligotrophic waters
at Lago di Monterosi, Italy (Goulden 1970), Wildcat
Lake, Washington (Davis et al. 1977) and Potato Lake,
Arizona (Whiteside 19 65). At Lake George in Australia,
Singh et al. (1981) found that elevated levels of
Pediastrum coinci ded with prolonged deep water (at
least 7 m) and at Lake Leake in Australia, Dodson
(1974a) found higher frequencies of Pediastrum asso -
ciated with ri sing lak e levels in the ea rly Holocene.
Pediastrum occurred in the LTC1 record during MIS 5.1
to early MIS 3 during high lake levels indicating clear,
oligotrophic waters (zone 7, Figure 2).
Remarks. Fossil zygospores and aplanospores (asexual
resting spores) of some of the eighteen genera of this
group of unbranched filamentous green algae have
long been recognised as common in microfossil
material prepared for pollen analysis (Davis 1975;
van Geel 1976, 1979). Zygnemataceae produce their
spores in spring in shallow (often less than 0.5 m deep),
stagnant, clean, oxygen rich warm water. They may
also occur near margins of lakes, in flowing water and
in moist soils or bogs (Transeau 1938, 1951; Grenfell
1995; van Geel and Grenfell 1996). The three taxa of
Zygnemataceae identified in this work were Debarya
glyptosperma, Spirogyra and Zygnema type.
Debarya glyptosperma (Plate 1, figure 4)
Description. Cells cylindrical to ovate, length 16–50 mm
and width 22–72 mm. Carinate with concave slope
between median and lateral keels, Corrugations run
radially with alternating ribs and furrows. Ribs usually
broader than furrows. Polar surface displays polar
Remarks. Early work in Western Australia identified
fossilised zygospores of Debarya (Transeau 1925) from
the Permian as Peltacystia (Balme and Segroves 1966;
Segroves 1967). Although a rare alga today, zygos-
pores of Debarya glyptosperma
(De Bary 1858) are so
characteristic that identification of this species is
reliable. Debarya glyptosperma found in archaeological
samples from a medieval settlement near Kootwijk, the
Netherlands and in palynological samples from sandy
subsoil of a bog on the Wietmarscher Moor, Germany
were useful indicators of the temporarily shallow
inundation of soils, either seasonal or during transition
to swamp (Ellis-Adam and van Geel 1978). Debarya
glyptosperma occurred in the LTC1 record in low
numbers at the end of MIS 5.1 (base of zone 7, Figure
2) and together with Anthoceros type indicated regular
incoming freshwater. Debarya glyptosperma also oc-
curred throughout the Holocene in all lake records
(zones 2 and 1 in Figure 2; zone 2 in Figure 3 and zone
4–2 in Figure 4) and, together with increased Glomus,
consistent values for Cyperaceae and the presence of
Typha, indicated lowered lake levels and increased
sediment infill at the lakes’ edges. This interpretation is
consistent with the formation of Fiery Creek swamp
at Lake Turangmoroke at *4300 calyr BP (*3900
C yr BP) identified by Crowley an d Kershaw (1994)
and a regional decline in precipitation recorded at
many sites in southeastern Australia between *2000
and *5000
C BP (Bowler 1976; Jones et al. 1998).
Spirogyra (Plate 1, figure 5)
Description. Spores ov al, radially symmetrical, length
20–80 mm. Wall shows longitudinal furrow.
Remarks. Spirogyra (Link 1820) occurs as mats of
green filaments in clear, oxygen rich, seasonally
fluctuating freshwater ponds, streams and lakes.
Hoshaw (1968) recorded optimal conditions for the
growth of most species in waters above 208C, indica-
ting stagnant, shallow lakes that more easily reach this
temperature. To reproduce, the cells of Spirogyra
conjugate in a manner similar to that of many fungi;
one cell empties content into another and the fusion
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produces a thick walled zygote, a resting stage for the
organism. The occurrence of Spirogy ra was most
prevalent through ou t the later parts of LTC2 and LB
records (zones 2 and 1, Figures 3 and 4) and, although
present sporadic ally an d/or in low numb ers throughout
LTC1 record, it was mostly observed in LTC1 in MIS 2
and the European phase (zones 4 an d 1). It often co-
occurred with Zyg nema type indicating freshwater
inundations into la kes that were beco ming more salin e
as they bec ame more stagnant and shal low (e.g. toward
the end of MIS 3 in the LTC1 record; zon e 5, Figure 2).
Zygnema type (Plate 1, figure 6)
Description. Hyaline, spheroidal spore length 35–53 mm
and 22–32 mm in diameter with hyaline, pitted surfaces.
Each pit is *2 mmdeepand*5 mmindiameter.
Remarks. Zygnema (Agardh 1824) is most profuse in
stagnant waters less than *0.5 m deep that warm
quickly because the optimum growthofthetaxonoccurs
at 15– 208C (Whitford and Schumacher 1963). Zygnema
is usually found in mesotrophic to eutrophic open
freshwater as green clumps and floating mats. Twenty
Zygnema species have been reported from Australia
(Entwistle et al. 1997). Here referred to a s Zygnema type
because other Zygnemataceous genera produce similar
spores. In this study, the representation of Zygnema type
throughout the LTC1 (Figure 2) and Lake Bolac records
(Figure 4) was consistent with its ecological profile.
4.3.2. Cyanobacteria (formerly blue-green algae)
(Plate 1, figure 7)
Description. Tubular hyaline sheaths irregularly cy-
lindrical, straight or slightly curved, length 60–250 mm,
width 15.5–32.5 mm, diameter 7–20 mm. Walls 1.5–
32.5 mm thick, thicker ones display lamellate structure.
Proximal end always rounded with pore 2–3 mmin
diameter. Distal end always open. Some contain
akinete (resting spore) with *1 mm pore at both ends.
Remarks. Se veral genera of the cyanobacteria have
sheaths on some of their cells that fossilise (e.g.
Gloeotrichia, Rivularia) (Livingstone and Jaworski
1980; van Geel et al. 1983, 1986, 1996). These
photosynthetic bacteria are commonly associated
with shallow, warm waters containing relatively low
nitrate (available nitrogen) as a result of their ability to
convert atmospheric nitrogen gas into nitrate (Scagel
et al. 1965; Jones 1994; Graham and Wilcox 2000).
They are also known to be predominant in eutrophied
systems (Hutchinson 1957) but decline when hyper-
trophic and higher turbidity levels are reached owing
to a lack of energy (light) available to drive the
nitrogen fixation process (Zevenboom and Mur 1980).
Van Geel et al. (1984, 1989) found that Gloeotrichia
was of great importance at a Late Glacial type section
at Usselo, the Netherlands, where, by undertaking
nitrogen fixation in pools of the earliest phase of
the Late Glacial, it made conditions suitable for the re-
establishment of other aquatic vegetation. The authors
also report from modern observation of the taxon that,
each year during late summer at a sandy pool dug in
a Dutch coastal dune area, G. echinu lata plays a
pioneering role in re-establishing aquatic plant life.
In the LTC1 record, increased Gloeotrichia num-
bers indica ted reduced lake levels consistent with a
decline in woody taxa (except Cupressaceae which
prospers in drier environments) and a return to drier
conditions during MIS 5.1–MIS 3 (zone 7 in Figure 2).
The pioneering role that Gloeotrichia played in fixing
nitrogen for plant growth can clearly be seen during
MIS 3 as peaks in this taxon at the bases of the empty
lake stages, indicated by low pollen sections between
zones 7 and 6 and 6 and 5 in Figure 2. During MIS 2,
Gloeotrichia occurred in the LTC1 record in substan-
tial numbers (zone 4, Figure 2) again indicating
decreased availability of nitrogen and accounting for
the depressed values for aquatic vegetation.
4.3.3. Bryophyta (liverworts, hornworts and mosses)
Bryophyte type spores (Plate 1, figure 8)
Description. Spor es circular, 32–36 mm in diameter.
Surface sculpture displays reticulate pattern.
Remarks. The bryophyte spore type identified occurred
in low levels throughout the Holocene in all records.
In broad terms it was representative of moist, shady
environments adjacent to the lake.
Anthoceros type (Plate 1, figure 9)
Description. Spores circular to sub-triangular, 45–
90 mm in diameter. Verrucate surface sculpture with
trilete mark.
Remarks. The class Anthocerotae (the ‘hornworts’)
possess characteristics of both liverworts and mosses.
Anthoceros is unique among the bryophytes because
spore production is a continuous process rather than
simultaneous with shedding only (Weier et al. 1970).
Anthoceros is widely distributed, and frequently found
on moist soils, in fields, on the surface of ponds and
ditches and as a pioneer alongside open water (Jahns
1980; Allaby 1992). Considered together with changes
in aquatic taxa, Anthoceros has been demonstrated as a
useful indicator of swamp development in southeastern
Australia (Dodson 1974a, 1974b; Dodson and Wilson
1975). Here recorded as Anthoceros type because
spores recorded also resem ble those of Lycopodiella
which are also present in the region. Lycopodiella is
similar in ecology to Anthoceros, growing among
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shrubs and sedges in swamps, in wet areas in soils or
on banks or slopes near streams (McCarthy 1998).
Anthoceros type representati on is greatest in the LTC1
record in MIS 5.1 (zone 7, Figure 2) and in all records
during the Holocene and in the European phase (zones
3–1, Figures 2–4), periods that were warmer and wetter
and during which swamps developed.
4.3.4. Mycota (fungi)
Remarks. Fungal spores that preserve as fossils are
dominated by ascospores and conidia (produced by the
Ascomycetes) and chlamydospores (Dematiaceae). The
abundance (and diversity) of fungal types is greatest in
peat deposits and much rarer in lake deposits, owing to
local production and limited dispersal of the spores.
Furthermore, there is a bias toward heavier fungal
spores with thick walls because many thin walled
spores do not fossilise (van Geel 2001; van Geel et al.
2003). Comparison of fossil spores with those from
extant genera is difficul t because reference collections
mainly focus on forms associated with crop plants and
there is little de tailed work by mycologists of fungi in
sediments (Jarzen and Elsik 1986). Unable to make
their own food by photosynthesis, fungi rely on host
plants or animals and identifications of fossilised
fungal spores can therefore enhance determinations
of vegetation and vegetational change by providing
specific information on their hosts, either living plants
(if fungi are parasitic), or their remains (if saprophy-
tic), the local presence of an imal dung, erosion and
burning (e.g. Davis et al. 1984; va n Geel et al. 2007).
Glomus (Plate 1, figure 10)
Description. Globose chlamydospores, aseptate and
inaperaturate. 17.5–138 mm in diameter, exclusive of
hyphal attachment. Wall thickness 0.5–5 mm, surfa ce
psilate to very finely scabrate.
Remarks. The endomycorrhizal fungus Glomus occurs
on a variety of host plants, and chlamydospores are of
regular occurrence on pollen slides. Anderson et al.
(1984) found G. fasciculatum in postglacial sediments
in Maine, USA, established with tundra vegetation on
new soils after the melting of Wisconsin ice. Glomus
chlamydospores form below the soil surface and are
not normally transported. Their increase during the
late glacial indicated increased soil erosion, and decline
during the Holocene indicated increased soil stability.
Glomus is present in Australia, is identified easily and is
a useful line of evidence in palaeoecology. Glomus was
a useful indicator of soil erosion in the catchments
of the two lakes studied. It occurred in highest levels
in the European zones of the records, likely reflecting
increased sediment loads to the lake as a result of
widespread clearing indicated by reductions in native
tree taxa (zone 1, Figures 2–4).
Neurospora crassa (Plate 1, figure 11)
Description. Brown, ellipsoidal, non-septate spore with
two protruding apical pores, each *1 mm wide. Size
22–30 mm 6 15–18 mm. Wall has fine longitudinal
Remarks. The asco spores of the saprotrophic fungus
Neurospora crassa germinate on rotting vegetable matter
after charring (Dennis 1968). The sterile environment,
rich in plant nutrients, toge ther with heat and chemical
by-products of the fire, favours ascospore generation
(Jacobson et al. 2004). Neurospora crassa asco spores
have been found in a layer of charred Molinia remains in
the Holocene bog Engbertsdijksveen (van Geel 1978),
were associated with high charcoal levels at a late
Holocene se ction from ‘Het Ilperveld’ in the Netherla nds
(Bakker an d van Smeerdijk 19 82) and were correl ated
with higher micro-charcoal frequencies in peat profiles
from North York Moors, northeast England (Innes
et al. 2004 ). In the US A, the species is a primary
coloniser of trees killed by wildfires including cotton-
wood stands in Rio Gran de and moun tain fore sts in
New Mexico (Jacobson et al. 2004). In association with
elevated charcoal levels, Neurospora crassa ascospores
clearly indicate fire in local proximity to the site and
contemporan eous ch arcoal production, pollen rain and
deposition of lake sediments. Neurospora crassa ascos-
pores have not previously been recorded in lake records.
In the study of Lakes Turangmoroke and Bolac , the
occurren ce of Neur ospora cra ssa gen erally coincided
with peaks in charco al indicating that much charcoal
was deri ved from so urces local to the sites. Peak s in the
charcoal, together with overall highest values for
Neurospora crassa in both the LTC1 and LB reco rds
at the commencement of the European period reflected a
change from low intensity, freq uent firing used by
Aboriginal people, to less fr equent and higher intensity
fires used to clear native vegetation duri ng early
European settlement (Cook 200 9 ).
Podospora type (Plate 1, figure 12)
Description. Ellipsoidal, smooth, dark brown ascos-
pores, 29– 63 mm 6 16–40 m
m. One pore, 2 mmin
diameter with annulus, protruding from directly be low
the apex. Basal end is bluntly conical.
Remarks. Ascos pores of the coprophilous Podospora
type have been recorded in low frequencies in samples
from archaeological sites where high densities of
megafauna or humans were present (van Geel et al.
1981, 1983; Buurman et al. 1995). Podospora type has
been found in deposits containing remains of
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Mammuthus in the Nethe rlands during the Moer-
shoofd Interstadial (Aptroot and van Geel 2006; van
Geel and Aptroot 2006), in Norwegian Calluna-
Sphagnum peat deposits with palynological evidence
for grazing (personal communication of BvG), in a
Roman Iron Age site in the Netherlands (van Geel
2001) and in late Holocene deposits from Mexico
(Almeida-Lenero et al. 2005). Most recently, van Geel
et al. (2010) recorded hundreds of Podospora conica
fruiting bodies, complete with ascospores, in late
glacial mammoth faeces from Cape Blossom, Alaska.
The presence of the intact fruiting bodies in the dung
ball provided evidence of coprophagy, a process that
may have been beneficial to mammoths in the way it is
for elephants today (van Geel et al. 2010). In the Lake
Turangmoroke record, the greatest representation of
Podospora type was during the European phase
(zone 1, Figure 2) consistent with the introduction of
sheep and cattle to the land around the lakes during
historical times. There are no Podospora recorded in
the Lake Bolac record, most likely because Lake Bolac
is much larger than Lake Turangmoroke, and deposi-
tion of fungal spores local in dispersal favoured the
smaller basin.
Sporormiella type (Plate 1, figure 13)
Description. Three to multi septate ascospores. Sig-
moid germinal aperture extends entire length of cell.
Ascospores break into separat e cells when ripe. Cel ls
12–28 mm 6 9–15 mm present as two types; end cells
show one flattened and one round, middle cells show
two flattened ends.
Remarks. Spores of Sporormiella are commonly
produced on the dung of herbivores such as cows,
sheep, goats, horses, rabbits, moose, deer, elk and
caribou (Davis 1975, 1976; Davis et al. 1977;
Pirozynski et al. 1988; Bell 1993, 2005; Davis and
Shafer 2006; Raper and Bush, 2009). The presence of
the fruiting bodies indicates the local presence of large
herbivores at a site. When found as fossils, Spor-
ormiella spores are a reliable proxy for the presence of
megafaunal biomass, their representation declining
during mass extinctions and increasing during the
introduction of domesticated livestock in recent
centuries (Davis 1987; Davis and Moratto 1988;
Burney et al. 2003; Robinson et al. 2005). Give n its
remarkable use as a marker determining faunal change,
the study of Sporormiella is especi ally use ful in
palaeoecological analysis. Van Geel et al. (2008)
recorded complet e fruit-bodies of Sporormiella in the
intestinal tract of the Yukagir Mammoth from norther n
Yakutia, Russia as eviden ce of coprophagy. In investi-
gating the development of late glacial plant communities
in North Ame rica with no modern an alogs, Gill et al.
(2009) used Spor ormiella spores to establish that
megafaunal decline closely pre ceded enhan ced fire
regimes that, together with decreased herbivory pres-
sure, led to the nove l plant co mmunities.
Van Geel (2001) noted that because representatives
of the related, also coprophilous genus Sporormia
produce morphologically similar spores that may
develop a sigmoid germinal aperture during germina-
tion (see Ahmed and Cain 1972), it is not possible to
differentiate fossil spores of these genera, and micro-
fossils should be referred to as Sporormiella type’. In
the Lake Turangmoroke record, occurrences of
Sporormiella type were greatest in the European phase
(zone 1, Figu re 2) with a significant peak in the LTC1
and Lake Bolac records during the latest Holocene,
However, at Lak e Bolac there is no peak in the
European phase after the introduction of domesticated
livestock, the reason for which remains unclear. Cook
(2009) reaso ned that the isolated Holocene repres enta-
tion was the result of a period of substantial but short-
lived runoff delivering these spores to the lake, not
normally reached given the large size of the lake.
Tetraploa (Plate 1, figure 14)
Description. Conidia verrucose of 3–4 columns of 2–4
cells, each ending in a septate appendage.
Remarks. The natural habitat of the fungus Tetrapl oa
includes leaf bases and stems just above the soil surface
of many herbaceous plants and trees (Dix and Webster
1995). At a site in the southwest of the United States,
abundant levels of Tetraploa have been found in historic
sediments due to the increased abundance of sen escent
plant material in freshwater marshes (Davis 1995).
Tetraploa was identified in the LTC1 and LTC2 records
4.3.5. Chironomidae (midges) (see identification guide
by Cranston (2000))
Description. Mandibles/labia (found in this work) 50–
75 mm long, 25–50 mmwide.
Remarks. Popularly referred to as ‘gnats’ or ‘midges’,
chironomids are among the most frequently collected
insects. Nearly all have aquatic larval and pupal stages
most of which remain fastened to submerged materials
until they rise to the surface to emerge as adults.
Heavily chitinised head capsules of larvae are abun-
dant in lake sediments, preserve well and occur in
waters from fresh to saline and fast flowing to still
(Williams 1980; Walker 1987, 1995). Fossil chirono-
mids have long been used to dedu ce palaeoclimates
(e.g. Anderson 1943) and detect human impact (e.g
Rodback 1970; Warwick 1975). Many species are
stenothermic, making them very specific climatic
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indicators and leading to the development and regular
use of chironomid-climate transfer functions (e.g.
Levesque et al. 1993, 1997; Walker et al. 1997;
Battarbee 2000; Brooks and Birks 2000; Wick et al.
2003; Larocque-Tobler et al. 2011).
In Australia over 50 genera and 130 specie s of
Chironomidae (Cranston 2000) have long been studied
to provide evidence of lake salinity levels. Examples
include pioneering work undertaken at Lake Gnotuk,
Victoria by Bayly and Williams (1966) and exploration
of Lake Werowrap, Victoria by Paterson and Walker
(1974) and Lake Keilambete, Victoria (see Kokkinn
1986). Recently Chironomids have been studied in
parallel with pollen analysis at Lake Barrine in
Queensland (Dimitriadis and Cranston 2001). Faunal
changes were correlated with fluctuations in water
levels, sediment influx, organic material, oxygen
availability and nutrient status mediated by changes
in the local climate and a shift from dry sclerophyll
vegetation to rainforest Dimitriadis and Cranston
2001). In the presen t study ‘Chironomi dae mandibles’
consistent with fossils recorded by van Geel et al. (1989)
were recorded during warmer and wetter environments
in the Holocene (zones 3–1, Figures 2–4).
4.3.6. Cladocera (water fleas) (Plate 2, figure 1)
Description. Spongy structure around ephippium of
Bosmina meridionalis (found in this work) measures
100–120 in diameter.
Remarks. These minute aquatic crustaceans are domi-
nant among zooplankton that preserve in lake
sediments. Common skeletal remains include parts of
the carapace (bivalve body covering behind the head),
ephippia (thickenings of the brood chamber), head
shield and appendages. Presently, *400 species of
cladocera are known worldwide; most inhabit fresh-
waters, only several marine species are known (Brusca
and Brusca 2003). Each species has different ecological
and climatic preferences making them sensitive recor-
ders of lake trophic status. Thus, fossil remains of
cladocera from freshwater lakes are widely used
proxies for the nutrient status of water bodies (e.g.
Birks and Birks 1980; Szeroczynska 2002) and transfer
functions are used to reconstruct climatic parameters
from cladocera assemblages (Lotter e t al. 1997, 2000).
Over 160 species of Cladocera have been described
for Australia from 50 genera with the majority found
in the families Daphniidae, Macrothricidae and
Chydoridae and smaller numbers in Sididae, Moinidae
and Bosminidae (Smirnov 1974; Smirnov and Timms
1983; Shiel and Dickson 1995). Their presence in
Australian salt lakes is restricted to a few species,
usually dominated by Daphniopsis pusilla (De Deckker
1982, 1988; Shiel et al. 1982; Williams 1986). In this
study, spongy material from around an ephippium was
identified as originating from Bosmina. Two genera are
recognised from Bosminidae; Bosmina (Baird 1845)
and Bosminopsis (Richard 1895). One species from
each is known in Australia; Bosmi nopsis dietersi
(Richard 1895), tropical in dist ribution, and Bosmina
meridionalis (Sars 1904), commonly found in south-
eastern Australia (Shiel 1995), making it reasonable to
identify the material observed as B. meridionalis.This
taxon is planktonic in the limnetic zo ne an d increa sed
levels are as sociated with increased nutrient levels in still
or slowly flowing wat er (Whiteside and Harmsworth
1967; Whiteside and Swindoll 1988). It was present
through the Holoce ne in the LTC1 and LB records.
4.3.7. Rotifera (wheel animalcules)
Remarks. Rotifers are microscopic (usually 42mmin
length) metazoa found in many freshwater environ-
ments, moist soil, on mosses and lichens and can even
grow on freshwater crustaceans and aquatic insect
larvae (Cuvier 1798; Friday and Ingram 1985;
Nogrady et al. 1993). Many Australian genera are
cosmopolitan (Shiel and Koste 1979; Koste and Shiel
1980; Ford 1982). Many species of rotifera produce
resting eggs (encysted embryos), some of which preserve
for many years withstanding adverse conditions and
enabling species to survi ve. They have been recorded in
sediment sequences from New Zealand (Duggan et al.
2002) but had not been rec orded in Australian
palaeoecological rec ords until this st udy (R. Shiel,
personal communication, 2004). Resting eggs of three
genera were identified as described in the following.
Brachionus (Plate 2, figure 2)
Description. Resting eggs ellipsoidal to ovoid, 158–
161 mm 6 95–98 mm, often open at one end. Wall is
reticulate and heterobrochate.
Remarks. The genus Brachionu s (Pallas 1766) (Brach io-
nidae; Wesenberg-Lund 1899) has 26 species in Australia,
many of which live in salt lake environments (Shiel 1995).
Brachionus plicatilis dominates halophytic lakes of
western Victoria (Williams 1978; Shiel and Koste 1986)
although there is evidence to suggest that too high a level
of salinity, such as that of seawater (38% at 308C), as
well as complete darkness and low temperatures blocks
hatching (Minkoff et al. 1983; Pourriot and Snell 1983).
Brachionus resting eggs occurred during MIS2 and in the
European phase of the LTC1 record (zones 4 and 1,
respectively, Figure 2) with rising levels of the submerged
aquatic Lepilaena cylindrocarpa type, tolerant of salinities
up to 27%, and an absence of fresh water Myriophyllu m,
indicating reduced lake levels and increasingly high
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Filinia (Plate 2, figure 3)
Description. Resting eggs globose, 58–100 mm 6 40–
95 mm, a furrow often encircles the body. Wall has
many bladders, each 1 mm diameter, large bladders are
10–18 mm diameter.
Remarks. Filinia species (Filiniidae; Bory de Saint-
Vincent 1824) are common among the planktonic
rotifers, usually found in fresh-brackish ponds and
lakes. Increased human impact around lakes has a
negative effect on egg numbers impairing their
production or preservation (Mu
ller 1970; Ruttner-
Kolisko 1972; Voigt and Koste 1978). In Australia the
genus contains *10 species, six of which also occur in
New Zealand (Shiel 1995). Filinia resting eggs occurred
in low numbers sporadically in the Lake Turangmor-
oke records (Figures 2 and 3). Their presence in the
Lake Bolac record (Figure 4), however, was generally
correlated with the presence of Botryococcus through-
out the Holocene until their disappearance in the
European phase, coincident with the timing of
Plate 2. Non-pollen palynomorphs identified (continued). Core and sample depths for specimens shown indicated in brackets:
LTC1 refers to Lake Turangmoroke core 1, LTC2 refers to Lake Turangmoroke core 2, LB refers to Lake Bolac. Scale bar
represents 20 mm. Figure 1. Spongy structure around ephippium of Bosmina meridionalis (LB 121–122 cm). Figure 2. Brachionus
resting egg (LTC1 1–2 cm). Figure 3. Filinia resting egg (LB 41–42 cm). Figure 4. Keratella resting egg (LB 41–42 cm). Figure 5.
Trichocerca, fragment of resting egg (LTC1 1–2 cm). Figure 6. Sponge spicule (LB 41–42 cm). Figure 7. Dinoflagellate cyst
(LTC1 1–2 cm). Figure 8. Foraminifera lining of test (LTC1 1–2 cm). Figure 9. Charred Poaceae epidermis (LTC2 67–68 cm).
168 E.J. Cook et al.
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widespread clearance of native vegetation, increased
erosion in the catchment and sedimentation in the lake.
Keratella (Plate 2, figure 4)
Remarks. The genus Keratella (Bory de Saint-Vincent
1822) (Brachionidae; Wesenberg-Lund 1899) has 13
species in Australia (Shiel 1995), commonly found in
billabongs and shallow wetlands with a minor presence
in reservoirs. In general, the genus Ker atella prefers
mesotrophic conditions where adequate food levels are
available. Keratella was limited in occurrence to parts
of MIS 2 and 1 in the Lake Turangmoroke records and
the Holocene in the Lake Bolac record (zones 2 and 1,
Figures 2–4) and presented in low numbers generally
consistent with the ecological preferences noted in the
Trichocerca (Plate 2, figure 5)
Description. Size 128–135 mm 6 80–116 mm, surface
texture has unevenly distributed sculpture.
Remarks. A variety of species of Tri chocerca (de
Lamarck 18 01) have bee n recorded in sal ine lake s in
western Victo ria (Timms 1981), Queensland (Timms
1987, 1998) and Western Australia (Brock and Shiel
1983). In Ger many, Mu
ller (1970) found that the resting
eggs of plan ktonic Trichocerca cylindrical disappeared
from the sediments of the Otterstedter Lake during
historical times, coinci dent with human impact recorded
in the pollen assemblage. Additionally, van Geel
(unpublished data ) linked reduced re presentation of
this taxon to human impact during historical times at
Lake Gos
z, Polan d. These studies show that Tricho-
cerca are excellent indicator taxa if identification is
pursued to species level. Trichoc erca occurred in low
numbers sporadically in the LTC1 record (Figure 2).
4.3.8. Porifera (sponges) (Plate 2, figure 6)
Description. Spicules *250–360 mm long, needle-like
in appearance.
Remarks. The body of a sponge is composed of a loose
aggregation of cells forming a hollow structure but
lacking intercellular nervous coordination. The body is
supported by an internal skeleton made from calcar-
eous or siliceous spicules or protein fibres (sporangin)
(Abercrombie et al. 1981; Martin 1983). Although
most sponges live in marine environments, *100
species are known from freshwater environments, the
majority of which are in the family Spongillidae. In
Australia, early estimates identified more than 10
genera and 24 species (Williams 1980); however, they
are rarely seen in salt lakes (De Deckker 1988). Sponge
spicules were most prevalent at the late glacial–
Holocene transition in LTC1 (zone 4, Figure 2), mid-
late Holocene in LTC2 (zones 3 and 2, Figure 3) and
very late Holocene and the European phase in LB
(zones 2 and 1, Figure 4).
4.3.9. Dinophyceae (Dinoflagellates) (Plate 2, figure 7)
Description. Cysts globose, hyaline, 40–58 mm ‘body’
diameter, appendages 19.5 mm long, 1–2 mm wide,
*5 mm apart.
Remarks. Dinoflage llates constitute a major componen t
of mar ine plankton and can also live in freshwater. They
reproduce by producing characteristic zoospores and,
during a resting stage, zygotic cysts (Beam and Himes
1980). Both the alga and cys t can be used in
palaeoenvironmental reconstruction (Mudie et al.
2004). In this study zygotic cysts occurred in the late
Holocene and the Europea n phase in the LTC1 record
(zones 2 and 1, Figure 2) and in the late Holoce ne in the
LTC2 record (zone 2, Figure 3). They are likely derived
from mater ial eroded from a local outcrop of marls,
laterite and limestone deposited during a major Tertiar y
marine transgression (Jackson et al. 1972; Jenkin 1988).
4.3.10. Foraminifera (forams) (Plate 2, figure 8)
Description. Test lining (found in this study) is
composed of a series of cells of increasing size. The
collection of cells measure 120 mm in length and
between 50 and 90 mm in width.
Remarks. Foraminifera construct architecturally com-
plex ‘tests’ (inner shells) made from calcium, chitin or
silica that are basic or many chambere d. Advanced
forams contain organic inner linings (membranes)
inside their tests as foundation for building and
protection against physical and chemical changes
Both tests and linings fossilise in sediments. Predomi-
nantly marine, foram species are be nthonic while a
small number are planktonic (Haynes 1981). A test
lining was fou nd in the European phase of the LTC1
record (zone 1, Figure 2) likely derived from erosi on
of local Tertiary marls, laterite and limestone.
4.3.11. Charred Poaceae epidermis (Plate 2, figure 10)
Description. Alternating long and short epidermal
cells, stomata clearly visible, usually 3–12 mmwide
and 2–3 times as long.
Remarks. The epidermis, or surficial layer of cells
covering a plant’s primary tissues, is perforated by
stomata, the function of which is to regulate gas
exchange between the plant and the atmosphere. These
pores are present on all aerial leaf and stem structures
as well as ovaries and anthers (Sinnot and Wilson
1955; Lowson 1962). Stomata occur on spore capsules
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of many Bryophyta but otherwise are restricted to the
vascular cryptograms and seed plants, and are
surrounded by a pair of guard cells that contain
chloroplasts (Robbins and Rick ett 1945; Weier et al.
1970). Morphologically it is possible to distinguish
between the microscopic remains of epidermal cells
(including stomata and guard cells) of different plant
types, even when charred or burnt and, where modern
reference material has been systematically compiled,
even within plant families where distinction below
family level on the basis of pollen morphology is
problematic (e.g. Poaceae, see Palmer 1976; Wooller
et al. 2003). In this study, increases in charred Poaceae
epidermis generally followed increases in charcoal in
all records (except during MIS 5.1–3 in the LTC1
record where there is only one representation) (zone 7,
Figure 2). Thus, it was possible to determine that
grasses were the major component of low level biomass
burning, principally during MIS 5.1 and the Holocene.
4.4. Description and illustration of unidentified
non-pollen palynomorphs
4.4.1. Types 180 and 209 (Plate 3, figures 1 and 2)
Remarks. Palynomorphs identified by van Geel et al.
(1989) as Type 180 and Type 209 were observed in this
study but remain unidentified. Type 180 (Plate 3, figure
1) has a distinctive hook shape, is hyaline and of
(greatest) diameter 43–125 mm. It is thought to be of
animal origin and was observed in a late Holocene
shallow pool deposit in the Netherlands by van Geel
et al. (1983). Type 180 was found in low numbers in the
European phase and later part of the Holocene during
saline lake conditions and/or swamp development in
all records presented here (Figures 2–4). Type 209
(Plate 3, figure 2) is a hyaline, isodiametric microfossil,
22–42 mm in diameter. The wall undulates to form 12–
26 protuberances (van Geel et al. 1989). It is likely
related to Type 66 which is correlated with meso-
trophic conditions (van Geel 1978) and Type 333 (van
Geel et al. 1981) which is found in the later part of the
Younger Dryas and at the commencement of peat
formation in the early Holocene in the Netherlands. It
is thought to be an algal cyst. In this study it is found
in greatest abundance during interstadials and stadials
when lake levels are low or variable. Here it is most
strongly associated with newly observed Type A3 in
LTC1 during MIS 5.2 (zone 8, Figure 2) and Type A9
in LTC1 during MIS 2 (zone 4, Figure 2).
4.4.2. Types A1–A12 (Plate 3, figures 3–14)
Remarks. Twelve palynomorph Types (A1–A12) re-
main unidentified. Type A1 (Plate 3, figure 3) is a
globose sp ore, 22–42 mm in diameter, exclusive of the
1.5–4 mm long and closely spaced protuberances from
the wall. In this study it is most strongly represented
in the European phase and the earlier part of the
Holocene of all records presented here (Figures 2–4)
during saline conditions and/or swamp development.
Type A2 (Plate 3, figure 4) is a pyramid shaped
palynomorph, 22 6 32 mm. It is four lobate and must
not be confused with Type 501 (Zopfiella lundqvistii;
van Geel et al. 1986; van Geel and Aptroot 2006)
which is a three lobate flattened fungal spore. Type A2
is perhaps an algal spore. Its representation is largely
restricted to parts of MIS 4 and 3 in LTC1 (zone 7,
Figure 2) and is associated with variable lake levels.
Type A3 (Plate 3, figure 5) is a dark brown triseptate
aporate ascospore, 10–12 6 26–30 mm. The axis of the
spore is curved with the outline convex on one side and
the other almost straight. Its greatest representation in
all records presented here (Figures 2–4) is during MIS
5.2 and the Holocene when lake levels are low.
Type A4 (Plate 3, figure 6) is a very dark brown
(almost opaque) monoseptate ascospore, 14–16 6 32–
34 mm exclusive of the 1–1.5 mm long very fine hairs on
the surface. Most often one cell is smaller than the
other. The spore is inaperaturate. Representation was
not sufficient to make correlations with other taxa.
Type A5 (Plate 3, figure 7) is a mid-light brown
monoseptate ascospore, 35–44 6 15–22 mm. It has
perforations at the apices. The axis of the spore is
curved with the outline convex on one side and the
other almost straight. It is most commonly represented
in Lake Turangmoroke during MIS 5.1 and the
Holocene (Figures 2 and 3). Type A6 (Plate 3, figure
8) is a dark brown cluster of cells of unknown origin.
The cells are globose, 12.5–25 mm in size. It is
inaperaturate. Representation was not sufficient to
make correlations with other taxa. Type A7 (Plate 3,
figure 9) is a dark brown triseptate ascospore 24–
28 mm in length. The spore is inaperaturate. Repre-
sentation was not suffici ent to make correlations with
other taxa.
Type A8 (Plate 3, figure 10) is a micro-sized piece of
wood with fungal growth. Pieces vary between * 30
and *70 mm in l ength. It occurred in Lake Turang-
moroke during parts of MIS 3 and the Holocene
(Figures 2 and 3). Type A9 (Plate 3, figure 11) is a thick
walled, globose microfossil, 26–47 mm in diameter. It is
perhaps an algal spore. It was found in all records
presented here (Figures 2–4) in the European phase
and later part of the Holocene during saline lake
conditions and/or swamp development. Its representa-
tion in the LTC1 record correlates with Types 209 and
A10 (Figure 2). Type A10 (Plate 3, fig ure 12) is a
zoological remain. It is similar in appearance to Type
509, identified as the armament of the pupal tergit of
170 E.J. Cook et al.
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the chironomid Glyptotendipes gr. pallens (van Geel
et al. 1986). It occurred in the European phase and
later part of the Holocene during saline lake conditions
and/or swamp development in all records presented
here (Figures 2–4). Type A11 (Plate 3, figure 13) is
an algal spore ovate in plan view. It has been noted in
other records from western Victoria dating from the
Tertiary to the Quaternary (B. Wagstaff, personal
communication, 2004). It occurred in Lake Turang-
moroke in MIS 3 and the Holocene (Figures 2 and 3).
Plate 3. Unidentified palynomorph Types observed. Types 180 and 209 were recorded by van Geel et al. (1983) and van Geel
et al. (1989) respectively, Types A1–A12 are newly identified. Core and sample depths for specimens shown indicated in brackets:
LTC1 refers to Lake Turangmoroke core 1, LTC2 refers to Lake Turangmoroke core 2, LB refers to Lake Bolac. Scale bar
represents 20 mm. Figure 1. Type 180 (LB 41–42 cm. Figure 2. Type 209 (a. LTC1 196–197 cm; b. LTC1 15–16 cm; c. LTC1 15–
16 cm). Figure 3. Type A1 (LTC1 117–118 cm). Figure 4. Type A2 (LTC1 196–197 cm). Figure 5. Type A3 (LTC1 49–50 cm).
Figure 6. Type A4 (LTC1 49–50 cm). Figure 7. Type A5 (LTC1 49–50 cm). Figure 8. Type A6 (LB 201–202 cm). Figure 9. Type
A7 (LTC2 60–61 cm). Figure 10. Type A8 (LTC1 45–46 cm). Figure 11. Type A9 (LTC1 201–202 cm). Figure 12. Type A10 (a–b.
LTC1 57–58 cm). Figure 13. Type A11 (LTC1 133–134 cm). Figure 14. Type A12 (LTC1 29–30 cm).
Palynology 171
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Type A12 (Plate 3, figure 14) is perhaps a mandible/
labia of an invertebrate. Its representation was not
sufficient to show on the diagrams but largely
coincided with remains of aquatic invertebrates.
5. Conclusions
An investigation of the diversity and indicator values
of NPP in two Late Quaternary lake records from
western Victoria, Australia, was undertaken. These
often overlooked microfossils contributed specific
additional information on past environments, human
impact and climate hist ory, to the general environ-
mental picture derived from pollen data. The results
have demonstrated that many of the NPP recorded are
specific algal, fungal, insect and other invertebrate
remains to which indicator values can be ascribed, and
that serve usefully as additional palaeoenvironmental
proxies, particularly for the refinement of past aquatic
conditions such as trophic stat us, salinity, depth and
temperature. The NPP have been described and
illustrated to exemplify the potential range of diversity
in Australian material. The study, and review, demon-
strates the usefulness of NPP in enhancing environ-
mental reconstructions in Australia, and elsewhere.
EJC gratefully acknowledges support from a Monash
Graduate Scholarship, Monash University Postgraduate
Research Travel Grant and Monash University School
of Geography and Environmental Science Travel and
Research Funding. Bert Roberts at the School of
Geosciences, The University of Wollongong, undertook
the OSL analyses with support from Australian Re-
search Council Discovery Grant A00104127 to A.P.
Kershaw and R.G. Roberts. Radiocarbon dating was
undertaken at ANSTO with funding from AINSE grant
97/196R and an AINSE Postgraduate Research Award
to EJC. We would like to thank Peter Kershaw and John
Grindrod for sharing their detailed knowledge of
Australian ecology and interest in the non-pollen
palynomorphs. The authors wish to thank P.F.M.
Coesel at the University of Amsterdam and M. Dingley
at the Royal Botanic Gardens, Sydney for making the
identification of Cosmarium distichum and for informa-
tion pertaining to its ecological preferences. J.N. Haas
and A.F. Lotter provided thoughtful comments on an
earlier version of this paper. We thank James B. Riding
and one anonymous reviewer for their very useful
suggestions for the improvement of the manuscript.
Author biographies
ELLYN J. COOK completed a BA Hons in soil
science then a PhD in Quaternary palynology at
Monash University in Australia, in which she explored
lunette-lake basins as sensitive recorders of environ-
mental change. Together with Sander van der Kaars,
she developed the first pollen-climate transfer func-
tions for use in Australia. Work in the INQUA
‘PALCOMM’ project focused on the southern con-
tinents and oceans led to publishing on glacial and
deglacial climatic patterns in the Australasian region.
Her interests include investigating patterns of inter-
glacial–glacial change and megafaunal extinction.
BAS VAN GEEL is Senior Lecturer in palaeoecology
and palaeoclimat ology at the University of Amsterdam.
During the last 40 year s he has focused on high
resolution Late Quaternary palynological studies of
lake deposits, fens, bogs and archaeological sites. He
leads research into
C wiggle-match dating of organic
deposits and understanding so lar forcing as a factor in
climate change as well as climatic teleconnections. In the
course of his palynological studies, he pionee red
investigation of the value of non-pollen palyn omorphs,
especially in combination with macr ofossil analysis.
During his work he has documented hundreds of such
‘Types’. Recently, he has applied these techniques to
ecological studies of mammoths and mastodons in
Siberia and North America.
SANDER VAN DER KAARS has worked on sites on
each of the continents as well as marine cores from
North America, Europe, Asia and Australia, ranging
in age from the Cretaceous to historical times. After a
Bachelors degree in Geology at the University of
Amsterdam, he moved to the Vrije Universiteit in
Amsterdam to work on the floral turnover at the K-T
boundary in Montana, Lower Tertiary coal deposits
from Colombia and the Neogene palynology of DSDP
site 603 in the NW Atlantic for a Masters in Geology.
He returned to the University of Amster dam to
undertake a Ph.D. in Quaternary palynology of the
Indo-Australian region using material from the Indo-
Dutch Snellius-II expedition. After studying sites in
Morocco, west Java, central Italy and the Adriatic, he
moved to Monash University and specialised in the
Quaternary history of the Indonesian-Australasian
region using marine as well as terrestrial cores. He
has produced over 50 records from the region and has
been engaged in debates over environmental change
and the arrival of people in Australia, as well as
investigating the effects of the Toba super-eruption.
JAN VAN ARKEL is a scientific photographer and
digital illustrator at the Institute for Biodiversity and
Ecosystem Dynamics at the University of Amsterdam.
He has always been interested in the nexus between art
and nature and studied monumental design at the
172 E.J. Cook et al.
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Rietveld Academie in Amsterdam, which he has used
to help him develop his skills in documenting nature in
pictures. Jan has worked across a range of scientific
subjects, from cataloging butterflies and phytoplank-
ton in zoology to illustrating the botanical remains
found in the intestines of mammoths.
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A general introduction to techniques which may be used to supplement pollen analysis. -K.Clayton
The fungal genus Neurospora has a distinguished history as a laboratory model in genetics and biochemistry. The most recent milestone in this history has been the sequencing of the genome of the best known species, N. crassa. The hope and promise of a complete genome sequence is a full understanding of the biology of the organism. Full understanding cannot be achieved, however, in the absence of fundamental knowledge of natural history. We report that species of Neurospora, heretofore thought to occur mainly in moist tropical and subtropical regions, are common primary colonizers of trees and shrubs killed by forest fires in western North America, in regions that are often cold and dry. Surveys in 36 forest-fire sites from New Mexico to Alaska yielded more than 500 cultures, 95% of which were the rarely collected N. discreta. Initial characterization of genotypes both within a site and on a single tree showed diversity consistent with sexual reproduction of N. discreta. These discoveries fill important gaps in knowledge of the distribution of members of the genus on both large and small spatial scales and provide the framework for future studies in new regions and microhabitats. The overall result is that population biology and genetics now can be combined, placing the genus Neurospora in a unique position to expand its role in experimental biology as a useful model organism for ecology, population genetics and evolution.
During 1974-1984, Lake Buchanan and seven peripheral pools usually contained water for only a few months each year, commencing in late summer. They ranged in salinity from 1 to 202 g 1-1, their waters were dominated by sodium chloride, but with Ca2+/Mg2+ ratios of c. 1, and were generally alkaline. The fauna of 53 species included three halobionts (e.g. Parartemia minuta, Diacypris compacta), 18 halophilics (e.g. Mytilocypris splendida, Trigonocypris globulosa, Microcyclops dengizicus) and many salt- tolerant freshwater forms, mainly insects. Overall, the fauna was distinctly Australian, but some prominent taxa found in southern salt lakes were absent and others were replaced by local endemics and tropical species. Past climatic cycles have probably influenced the composition of the fauna.