Proceedings of the

Linnean Society of New South Wales

Vegetation Patterns across the Sydney Basin during the
Last Glacial Maximum based on Plant Biogeography,
Ecology, Geomorphology and Climate
Doug Benson
Honorary Research Associate, Australian Institute of Botanical Science,
National Herbarium of New South Wales, Botanic Gardens and Domain
Trust, Sydney NSW 2000 Australia. [email protected]
ABSTRACT
Natural History in all its Branches Despite the importance of the Last Glacial Maximum (LGM) in the
formation of modern Australian vegetation, no comprehensive account
exists of the likely vegetation of this period in the Sydney Basin. Likely past
Benson, D. (2024). Vegetation LGM vegetation events and patterns for this region are proposed, based on
patterns across the Sydney geomorphological, palynological and archaeological studies, combined with
Basin during the Last Glacial current geographical distributions of endemic, restricted and common plant
Maximum based on plant species, inferences from basic ecological characteristics, observed vegetation
biogeography, ecology, community interactions, and morphological or genomic variation.
Based on ecological attributes (stress-tolerators, competitors, ruderals),
geomorphology and climate.
and geographic distributions, individual species are associated with likely
Proceedings of the Linnean past survival during the drier and colder LGM climate (28 000-18 000 years
Society of New South Wales 146, BP), across the different landscapes within the region. These include the old,
1-47. climatically buffered, infertile (OCBIL) Sydney sandstone plateau systems;
the old but more fertile shale (OCFEL) areas (Illawarra, western Sydney,
Published on 5 March 2024 at Central Tablelands); the younger fertile alluvial lowlands (YODFELs); and
https://openjournals.library.sydney. the now-inundated LGM continental shelf/coastal plain.
edu.au/index.php/LIN/index Potential LGM vegetation scenarios include Blue Mountains moisture-
related stress-tolerant heath, mallee, seepage and cliffline vegetation on an
Manuscript received 30 May LGM subalpine zone (>800 m elev.) with snowmelt and effectively better
2023, accepted for publication 6 moisture; dry grassland and mallee shrubland on the Central Tablelands and
December 2023 western Sydney shale areas; and eastward movement of western clay soil
species through Hunter Valley/Southern Highland connections and sclerophyll
Keywords: competitors, continental shrubs across mid-elevation dry sandstone. The importance of the coastal
shelf, Last Glacial Maximum plain/continental shelf, including coastal wallum vegetation, is suggested as
(LGM), OCBILs, stress-tolerators, the origin of many species’ distribution disjunctions.
vegetation floristics. By interweaving circumstantial evidence and speculation, hypotheses
and narratives can be developed to provide a spur to plant ecological and
biogeographic research and advance research in the field.

INTRODUCTION
Over the past half-century much effort has gone into mapping and
describing the vegetation communities of New South Wales, their patterns
in relation to soils and current climate, and their ecological responses to
impacting factors particularly fire. Rarely has current vegetation or work on
rare species been linked with possible past vegetation histories. There are
PO Box 291, Manly NSW 1655 exceptions mainly relevant to very distant time periods, e.g. highlighting
https://linneansocietynsw.org.au the antiquity of species such as the Wollemi Pine, Wollemia nobilis (from
[email protected] a 200 million-year-old plant family) (Jones et al. 1995), formation of
0490 542 524 Gondwana rainforests during the Cretaceous period (about 140 million
years ago), and evolutionary histories of various phylogenies over similar
ISSN 1839-7263
time periods (e.g. Weston and Jordon 2017; Bryceson et al. 2023).

Proc. Linn. Soc. N.S.W., 146, 2024 1

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

The Last Glacial Maximum (LGM) (28 000- Rare plants may tell us about their past
18 000 years BP) is an inherently interesting Some plant species are widespread and common
period, encompassing the Ice Age era in the across a landscape, some are widespread but rare,
Northern Hemisphere, popularised in commercial others are rare but locally common, or sometimes
entertainments, and occurring within the period very localised (Rabinowitz et al 1986; Kunin and
of Aboriginal occupation in eastern Australia Gaston 2012). Plant distributions are generally
(Attenbrow 2002). The Sydney Basin landscape is considered in relation to ecology and habitat;
a major Australian biogeographical area, but despite concurrent distributions may be generalised into
the influence of the severe climatic disruptions of plant communities and interpreted through landscape
the LGM on the formation of the modern vegetation and vegetation mapping. This classification process
(Mooney et al. 2017), there is no comprehensive depends mainly on the distributions of the widespread
account of its vegetation history in this period. The and common species. Rare and very localised species
relatively dry climate, the stability of the sandstone do not contribute much to the landscape picture,
landscapes and frequency of major fire events have left though conservation management is often focussed
few surviving sites with evidence of past vegetation on our inherent interest in them.
histories for geomorphological, palynological and Plants may be rare because we have destroyed their
archaeological studies. Peat-forming conditions are habitat (e.g. clearing for agriculture or urban development)
limited and most begin more recently than the LGM. or changed the conditions under which they prospered
The various historical sources, pollen, fossil (e.g. grazing by livestock or rabbits) (NSW Office of
testate amoebae, charcoal and the geomorphology Environment & Heritage 2023). However, some plant
only provide a broad, non-specific, picture of LGM species are naturally rare, that is, they have always been
structural vegetation structure of the Sydney area of limited occurrence in the current landscape, at least
during the LGM (e.g. Hesse et al. 2003; Robbie and as we understand it. Within the Sydney region many of
Martin 2007; Chalson and Martin 2008; Mooney et al. these rare species are geographically restricted, many
2021). There was vegetation, except perhaps in higher are restricted to sandstone habitats; many are Central
parts of the upper Blue Mountains. There is no evidence Coast/Central Tablelands endemics; and some species
of glaciation, though treeless conditions are inferred by occur naturally in very limited local clusters. Others
limited occurrences of clifftop sand dunes (Hesse et al. may have populations separated by large distances, i.e.
2003). There was some eucalypt-dominated vegetation disjunct distributions, in relation to the species’ apparent
in sheltered sites in the lower Blue Mountains (Robbie dispersal mechanisms, often on related sandstone
and Martin 2007), and essentially shrubby vegetation or similar landscapes, which are themselves widely
(i.e. with much less tree cover) on the coastal sandstone separated (particularly on the South Coast and Southern
plateaus. There was contrasting grassy but treeless Tablelands). Over long periods of time, small differences
vegetation on shale soils in western Sydney (Chalson between isolated populations of a species may develop
and Martin 2008). in response to differing local habitat/climatic conditions
Can the contemporary distributions of extant or from random drift, a process known as vicariance.
plant species, at least for those with particular Major differences may indicate the evolution of separate
ecological characteristics, expand this structural species over a long period of time. Slight morphological
picture? Biogeography relates occurrences of plant variation, recognised by botanists as subspecies,
species with landscape and climate at continental varieties or local forms, may be evidence of change over
and local scales; it has been integral to advances shorter or more recent time periods.
in scientific thinking on major issues including The Sydney area
evolutionary history and continental drift (Weston and The Sydney area here includes both the Sydney
Jordon 2017). Evolutionary studies use biogeographic Basin Bioregion, a major sedimentary geological
information to interpret taxa changes over millennia. basin, and the eastern edge of the South Eastern
Can contemporary plant distributions be interpreted Highlands Bioregion with contrasting metamorphic
as “fossil” evidence of past distributions? Can geology (NSW National Parks and Wildlife Service
current distributions of rare plants, unexpected 2003). The area covers the NSW Central Coast and
occurrences of more common ones and geographical Central Tablelands Botanical Subregions (Figure
distributional limits of others, together with their 1) (Benson and McDougall 1993), extending from
ecological characteristics (particularly distance Newcastle to Nowra and inland to Oberon and
dispersal and longevity), tell us something about past Bathurst. It includes major areas of National Park
vegetation patterns over more recent periods of time, (NP) including the one million ha of the Greater
and specifically during the Last Glacial Maximum? Blue Mountains World Heritage Area (GBMWHA).

2 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

lying Cumberland Plain, which includes

most of western Sydney and extends about
50 km inland, and includes the alluvial
flats of the Hawkesbury-Nepean River.
The Cumberland Plain is surrounded by
an arc of older Triassic Hawkesbury and
Narrabeen Group coastal and mid-elevation
sandstone plateaus (Figure 2), including
Brisbane Water, Ku-ring-gai Chase,
Royal, Nattai, Blue Mountains, Yengo and
Wollemi National Parks. Annual rainfall is
high on the coast (1200 mm pa), dropping
to about 600 mm in western Sydney, and
then increasing with elevation to about
1200 mm in the upper Blue Mountains.
Soil fertility in the Sydney Basin is
generally low, especially on the Triassic
sandstones, which are characterised by
shrubby sclerophyll vegetation structurally
dominated by eucalypt woodland and
open-forest (Keith 2004). Much of the
Figure 1. Sydney area showing Central Coast and Central Tablelands area is within the Greater Blue Mountains
Botanical subregions. World Heritage Area (GBMWHA), which
was listed for World Heritage in 2000 for
outstanding universal values related to the
exceptional richness and unique values of
its biodiversity (NSW National Parks and
Wildlife Service 1998).
Biodiversity
Sydney area vegetation has been of
scientific interest since Joseph Banks and
Daniel Solander made the first botanical
collections at Botany Bay in 1770 (Benson
and Eldershaw 2007). The new plants
stunned the scientific community in Europe
and further exploration of the flora of the
Blue Mountains did not disappoint. The
Sydney area includes about 2000 native
plant species, more than 100 of which
are endemic. The case for listing the Blue
Mountains for World Heritage cited the
biodiversity richness of the eucalypt forest
flora (almost 100 species), the diversity of
habitats, and the number of endemic species
Figure 2. Sydney area landforms, the darker green dissected (NSW National Parks and Wildlife Service
sandstone landscapes contrasting with the low elevation western
1998). The Sydney Montane Heaths, for
Sydney Cumberland Plain.
example, have some outstanding examples
The distance from the coast west to the upper Blue of vicariant biogeography, the phenomenon
Mountains is about 100 km with an elevation rise whereby closely related species replace one another at
from sea level at Sydney to about 1200 m in the Blue different locations (Keith 2004). Many species have
Mountains and Great Dividing Range. Geologically localised and/or disjunct distributions, for example,
the area is centred on the Triassic Wianamatta Shale Darwinia spp., Persoonia spp. and mallee eucalypts
and Tertiary alluvial sandy clay deposits of the low- (Rutherford et al. 2016).

Proc. Linn. Soc. N.S.W., 146, 2024 3

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

The high species diversity associated with Scope of this work

low nutrient landscapes worldwide, such as in the Research on climate change is providing
southwest Australian Floristic Region, the Greater increasing evidence of past Pleistocene and Holocene
Cape area in South Africa and Pantepui in South climates, previously only open to speculation, giving
America, has been recognised in the OCBIL theory. greater confidence for vegetation interpretation. This
This multi-hypothesis formulation was developed work develops an historical ecological narrative
to increase understanding of the evolution, ecology describing likely past vegetation patterns across the
and conservation of biological and cultural diversity Sydney Basin, using climatic, geomorphological,
on old, climatically buffered, infertile landscapes, palynological and archaeological studies, together with
designated OCBILs (Hopper 2021, Hopper et al. 2021). current geographical distributions of plant species,
Hopper has suggested that on OCBIL landscapes and inferences from basic ecological characteristics,
such as the Sydney Blue Mountains sandstone area, vegetation community interactions and morphological
sclerophyll flora has evolved with a particular range or genomic variation (together with a set of assumptions
of morphological and ecological characteristics, about past vegetation behaviour). The time frame
including stress-tolerance features, accentuated being considered is from the Pleistocene Last Glacial
persistence, commonly clonal old individuals, and Maximum (LGM) (28 000-18 000 years BP), through
slow individual growth. Other OCBIL ecological the subsequent warming events of the early and mid-
features include: limited dispersal ability; seed simply Holocene, to the onset of increased climatic variability
being shed around the parent resulting in fidelity to and the fluctuating climatic patterns associated with
certain habitats thereby increasing local endemism; increased frequency and/or strength of the El Nino
and nutritional and other specialisations such as cluster Southern Oscillation (ENSO). This first paper uses
roots, hemi-parasitism and ant-adapted dispersal plant species distributions and ecology to infer evidence
(exemplified in many Fabaceae - Berg 1975). Species of Last Glacial Maximum vegetation in the Sydney
with these stress- tolerator characteristics, including Basin. Vegetation areas dealt with include the coastal
both dicot shrubs and monocot geophytes make up sandstone plateaus and LGM continental shelf, upper
40-80% of local Sydney heath floras (Keith 2004), Blue Mountains and Tablelands, and Western Sydney
including many rare and endemic species confined to landscapes. Following work will cover the impacts of
plateaus and ridgetops. Some terrestrial orchids have the Holocene warming (14 000-6 000 years BP), and
particular soil, microbial and pollinator requirements the subsequent progression into the contemporary
restricting their habitats. Stress-tolerator and OCBIL period (the last 6 000 years) associated with increased
characteristics align closely with characteristics likely climatic variability and the development of the El Nino
to have facilitated species survival in situ during the southern oscillation (ENSO).
harsh conditions of the LGM.
Late Pleistocene climates in southeastern
Associated with the sandstone OCBIL landscapes
Australia
are more fertile clay soil landscapes exposed on
During the last 100 000 years of the Pleistocene
interleaved shales forming old, climatically-buffered
Period, conditions became increasingly colder and
relatively fertile landscapes (OCFELs) (Hopper et al.
drier in southeast Australia, with a progressive buildup
2021). Of broadly similar age to the OCBILs these
of ice (Mooney et al. 2017), but with enhanced
support grassy woodlands and rainforest species
effective precipitation during some intervals, such as
with some OCBIL characteristics indicative of long
cool and humid conditions from 57 000-29 000 years
periods of stable existence. In contrast, areas of
BP. The severe climatic conditions culminated in the
younger fertile alluvial and riparian soils, and Recent
Last Glacial Maximum (LGM) extending from about
dune soils, may be classed as YODFELs, i.e. younger,
28 000-18 000 years BP. Average global temperatures
often disturbed, more fertile lowland landscapes
were about 7 °C colder than in the 20th century. Sea level
(Hopper 2021), and may have quicker growing and
dropped to 125 m below that of today. Northern Europe
more readily-dispersed species than the OCBIL
and North America were covered in massive ice sheets.
sclerophylls. The importance here is the age of the
Britain was covered in ice, except for Graminoid and
landscape rather than just the fertility. Some classic
Forb Tundra in the southeast, to become Deciduous
OCBIL landscapes such as the Sydney sandstones
Forest in the modern era (Binney et al 2017).
have probably been in existence contemporaneously
Less severe climatic conditions occurred in
with mesic rainforest on OCFEL landscapes since
Australia, which had only minor periglacial activity
Gondwanan times. The juxtaposition of OCBIL,
(Kosciusko and Tasmania) but was essentially
OCFEL and YODFEL ecosystems is a major
free of ice and mostly remained vegetated, but
contributor to Sydney area habitat diversity.

4 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Figure 3. Sea level change over the last 150 000 years from Waelbroeck et al. (2002) (upper graph) with
glacial and interglacial periods highlighted (U3A) (lower graph).

LGM 28000-18000 yrs BP General Coast Montane Present day References

Williams et al.,
winter up to 8
Mean air Temperature 8-10 0C lower 12-20 0C 2009, Barrows et al.
C lower
0
2022
snow at high Williams et al.
Up to 50% lower, 700-1600 mm
Rainfall elevations 2009, Mooney et al.
pa, light snow
low evaporation 2021

Atmospheric CO2 185-190 ppm 280 ppm Mooney et al. 2017

Strong winds
Winds 4 times more Hesse et al. 2003
frequent
Southern Ocean temperatures -2 to -60 C lower 18-230 C Williams et al. 2009
125 m lower at 21-
Hawkesbury drowned Reeves etal 2013
Sea level 19,000 yrs BP;
river valley estuary Lewis et al 2013
8-15 km wide with low ?freshwater swamps
LGM Coastal plain sandstone plateaus and on Parramatta, Albani et al. 2015
podsolised dune sand Georges, Hacking
Table 1. Summary of major Last Glacial Maximum (28 000-18 000 yrs BP) climatic and landscape conditions
relevant to the Sydney area.

Proc. Linn. Soc. N.S.W., 146, 2024 5

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

experienced a period of high aridity, and was much landscape change peaking with the LGM was the
drier and windier, with increased evapotranspiration, development of a coastal Plain (Albani et al. 2015)
and cooler temperatures (Mooney et al. 2017). In adding an area of about 250 000-300 000 ha between
mainland southeast Australia LGM temperatures the Hunter and Shoalhaven Rivers and, in relative
were up to 8 °C lower than today and up to 4 °C terms, pushing the hinterland and mountains 8-15 km
cooler in Tasmania. Sea surface temperatures were further inland and the mountains another 150 m higher.
up to 5 °C cooler (current annual range 18-23 °C off
Increased warming and the end of LGM climatic
Sydney) with rapid warming after 20 000 years BP to
conditions
attain present values by 15 000 years BP (Williams et
By 17 000 BP post-LGM increases in temperature
al. 2009). Winter rainfall was much reduced although
were evident across Australia (Reeves et al. in Mooney
periodic extreme flood events are evident in the fluvial
2017). Low sediment rates in Sydney cores confirm
record (Williams et al. 2009). Atmospheric CO2 at
LGM low rainfall conditions continuing well into the
180 ppm was 25-30% lower than in the following
late Pleistocene (Mooney et al. 2021) with minimum
Holocene (270 ppm) (Mooney et al. 2017) and less
effective precipitation in southern Australia from 14
than half current levels of 417 ppm. Sea levels during
000 to 12 000 yrs BP (Williams et al 2009), delaying
the LGM were about 125 m lower than current pre-
the recovery of tree-dominated communities. There
industrial levels (Sloss et al. 2007, Lewis et al. 2013,
were relatively low levels of biomass burning in the
Reeves et al. 2013 (Figure 3) (Table 1).
period 24 000 to 18 000 years BP (Mooney et al.
In the period 28 000-18 000 years BP, areas of
2011). In Western Tasmania Cool temperate rainforest
southwest Western Australia that support eucalypt
developed again after about 14 000 yr BP, with
forests today appear to have been shrublands
maximum development of Nothofagus cunninghamii
(Sniderman et al. 2019). LGM vegetation in western
rainforest in the early Holocene (Colhoun 2000).
Tasmania was subalpine woodland and shrubland,
From 14 000 to 9 000 years BP (Pleistocene–
or alpine grassland, herbland, and heathland, with
Holocene transition) temperature increased
sclerophyll woodland and rainforest taxa below 100
considerably (approx. 4 °C). Warmer wetter conditions
m on coastal valley floors. Major vegetation changes
and low fire activity associated with climatic stability
were primarily climatically driven, responding
peaked about 6 000 years BP (Black et al. 2008; Moritz
to variations in temperature in the west and to
et al. 2009; Das et al. 2019). In contrast, from about 6
temperature and precipitation in the east (Colhoun
000 years BP, the El Niño phase of ENSO developed
and Shimeld 2012).
in frequency and/or strength, increasing climate
In south-eastern Australia late Pleistocene
variability, a significant influence on vegetation
paleochannels of Murray-Darling Basin rivers were
composition, with impacts felt through drought and
larger, with high seasonal discharge and sediment
fire (Mooney et al. 2017). As temperatures rose from
loads indicating increased moisture availability in
about 18 000 years BP, increasing ocean volumes led
Tableland catchments from snowpack, enhanced
into the early Holocene sea level rise of c. 60 m, from
orographic rainfall, or CO2 feedbacks with vegetation
about 11 000 – 7000 years BP during deglaciation
cover (Mueller et al. 2017, Hesse et al. 2018, Barrows
over most of the Earth (Smith et al. 2011). Sea levels
et al. 2022). Modelling of precipitation suggests LGM
temporarily reached more than a metre above current
snowfall was a common winter occurrence across
levels between 7 000 and 2 000 years BP before falling
southern Australia (Mueller et al. undated). Periglacial
back to current levels (Lewis et al 2013, Sloss et al.
conditions with mean winter temperature 8 °C colder
2007). The vegetation responses to these post-LGM
than present affecting much of southeastern Australia,
changes will be dealt with in later papers.
down to 680 m on the Southern Tablelands, are likely
(Barrows et al. 2022). Northern Tablelands LGM
METHODS
montane vegetation appears wetter than previously
thought, with higher moisture conditions and a more Plant ecology and postulating past vegetation
positive water balance than occurs today (Ellerton et al. patterns
2017). On either side of the LGM peak (24 000 years Sources of botanical evidence to reconstruct
BP), Willandra Lakes in western NSW retained water past vegetation patterns and species groupings
and supported human activity; elsewhere in Australia include current flora distribution patterns, ecological
during the cold and dry times, water was present to strategies and tolerances, and historical pollen and
sustain human populations (De Deckker et al. 2020). fossil evidence. Using this information, together with
While Sydney area vegetation changes were a set of assumptions about past vegetation behaviour,
primarily climate-driven, as in Tasmania, a substantial species groups have been retrofitted, as it were, into

6 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

a number of past physical landscape scenarios based influences are consistent with current
on currently accepted views of late Pleistocene- plant strategy schemes and knowledge,
Quaternary climatic conditions (Table 1). i.e. competitor/stress-tolerator/ruderal
In doing this the following ecological strategies, fire response categories, etc.
assumptions have been made: And the following provisional interpretations applied:
1. same landform/topography interactions 1. That for species with limited dispersal
(with additional LGM coastal plain), e.g. abilities (i.e. stress-tolerator/OCBIL
rainfall increases with elevation and declines characteristics) disjunct distributions are
with distance from coast, with rain-shadow likely to be evidence of previously more
effects as today; mean temperature declines extensive distributions, fragmented by
with elevation, i.e. rainfall and temperature subsequent unfavourable climatic or other
are related to geomorphology/elevation; conditions, rather than new colonisations.
2. current existing pool of south-eastern 2. That for species with readily animal-
Australian species is available, but may or bird-dispersed propagules, disjunct
have had different geographic distributions; distributions are more likely to be more
3. species niche conservatism ecologies recent introductions/colonisations, rather
constrained by current basic ecological than fragmentations.
envelopes, i.e. ecological limitations are Using National Herbarium of New South Wales
imposed by soils and geology though (PlantNET) and Australasian Virtual Herbarium
climatic envelope may be more flexible; (AVH) data sources, species occurrence maps were
4. 10-20 000 years is too short for species level examined to identify species with particular patterns
evolution, but allowing small genetic and/or of disjunction, with particular focus on endemics,
morphological differences (e.g. subspecies, dominants, edge of range, etc., that could indicate past
forms) to develop as species respond to landscape change, e.g. climatic isolation, loss of LGM
climate change through migration, isolation coastal plain, distance invasion, and any reported
and local extinction; evidence of taxonomic or genetic variation (e.g. Table
5. species interactions with past climate 2). Geographical areas or topographic sites with high

Life form
Pop. distr.
Habitat Species Distribution CSR/fire
type
response
Zieria covenyi Katoomba ST/Respr. single
Pultenaea genowlan Newnes Plateau ST/Respr. single
Eucalyptus camfieldii Ku-ring-gai to North Head ST/Respr. disjunct
Ridgetop sandstone Eucalyptus gregsoniana Newnes Plateau ST/Respr.
Melaleuca deanei ST/Respr. disjunct
Clarence, Mullion
Acacia meiantha ST/Respr. disjunct
Range, Turon
Micromyrtus minutiflora Castlereagh area Rud/fire sens disjunct
Dillwynia tenuifolia Castlereagh area Rud/fire sens disjunct
Low nutrient clay Pultenaea parviflora Castlereagh area ST/Respr. single
Allocasuarina glareicola Castlereagh area ST/Respr. single
Grevillea caleyi Terrey Hills Rud/fire sens single
Wianamatta Shale Pimelea spicata western Sydney, Kiama ST/Respr. disjunct
woodland Marsdenia viridiflora Mt Annan ST/Respr. disjunct
Asterolasia buxifolia Hartley Rud/fire sens single
Leptospermum deanei Lane Cove valley Rud/fire sens single
Riparian
Boronia deanei Newnes Pl. Kanangra ST/Respr. disjunct
Hibbertia hermanniifolia Bents Basin disjunct
La Perouse, Agnes
Coastal sand Banksia aemula ST/Respr. disjunct
Banks, Sth limit
Eucalyptus cunninghamii Bl Mtns, Wanganderry ST/Respr. Wide disjunct
Montane heath
Isopogon prostratus Newnes Pl.-Vict. ST/Respr. Wide disjunct
Ridgetop metasediments Eucalyptus pulverulenta Mt Blaxland, Bombala ST/Respr. Wide disjunct
Table 2. A selection of rare plant species, local endemics etc, illustrating life forms and types of distributions. ST/
Respr.=Stress tolerator/ resprouter.

Proc. Linn. Soc. N.S.W., 146, 2024 7

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

numbers of rare or endemic species or with clusters being fire ecology responses and, more recently,
of species with similar disjunction patterns were seed biology and genetics of rare and endangered
identified. species. Much of the ecological data applicable to
these lifeforms is taken from the Ecology of Sydney
Ecological strategies and tolerances of individual
Plants series (Benson and McDougall 1993-2005),
species
major ecological reviews including Fabaceae
Various groupings of plant species based
(Auld 1996), Myrtaceae (Myerscough 1998),
on functional traits have been developed to
Orchidaceae (Weston et al. 2005), Proteaceae
conceptualise ecological responses (Noble and
(Myerscough et al. 2006) and Rutaceae (Auld
Slatyer 1980; Keith et al. 2007). To consider the
2001) together with many personal observations
likely responses of individual plant species in past
made by the author over a long career in Sydney
climate scenarios, a combination of Grime’s (2006)
plant ecology.
Competitor/Stress-tolerant/Ruderal (CSR) lifeform
strategies (Grime 2006; Grime and Pierce 2012) and Historical pollen and fossil evidence
Australian fire response terminology (Gill 1981; Worldwide, pollen analyses are a primary source
Keith and Tozer 2012) is used (Table 3). Three broad for historical vegetation interpretation. Records provide
groups are recognised: stress-tolerators/resprouters an extensive late Quaternary vegetation history for
(many OCBIL species), ruderals/fire-sensitives Tasmania (Colhoun and Shimeld 2012), for example,
and competitors. Competitors are not generally but because of limited suitable deposition sites, there
recognised in fire ecology (though included in some is very little plant macro-fossil or pollen evidence
schemes (Keith et al. 2007) but some species with for this period for Sydney. References with pollen
both resprouter fire responses, and vigorous seeding core diagrams were studied for additional ecological
recruitment capabilities (facultative resprouters), signals from clearly defined taxa. Helene Martin and
such as many tree eucalypts (Nicolle 2006) are her students have produced important studies but
clearly competitive. It should be recognised that pollen identification is generally limited to genus or
characteristics such as the large size and quick family, limiting ecological inferences in a floristically
growth of competitors make them competitive in rich area. Chalson and Martin (1995) provide a pollen
today’s conditions though in dry, low CO2 LGM reference set for 20 of the important Myrtaceae
environments these characters are likely to have species, but the Sydney area has over 100 eucalypts
made them uncompetitive (Woillez et al. 2011). (Hager and Benson 2010) and about 30 Leptospermum
While collective and institutional efforts have species. Within a family or genus, individual species
historically been focused on formally describing may have very different growth-forms and ecologies
plant species, much less effort has been focussed but similar-looking pollen. Family Casuarinaceae
on describing their ecologies, the main exceptions taxa in the Sydney area include a large fire-sensitive

Grime CSR functional classes Australian fire ecology strategies Notes

Stress Tolerators Primary resprouters Long-lived sclerophyll or mesic
Small, long-lived, prevail in habitats Resprout from basal or epicormic trees, shrubs and graminoids,
with limited resources, low rates of shoots post- fire; generally low seedling generally slow-growing and low
photosynthetic activity and respiration recruitment success recruitment, often vegetative
spread, poor competitors
Ruderals Fire-sensitive/Obligate seeders Shortlived but includes early
Short-lived, have low leaf dry weight, Killed by fire; recruitment from seed maturing eucalypts such as
longer flowering period, high rate either soil-stored (with dormancy) or Eucalyptus oreades with
of photosynthetic capacity and canopy-stored (serotinous) exploiters of longevities up to 100 years
respiration, and high leaf nitrogen temporally available resources
content
Competitors Competitors not specifically recognised Competitors with both
High levels of canopy height, leaf dry in fire succession, but important in resprouting survival and
weight, and maximal lateral spread, directional rather than cyclic change, high seedling recruitment
rapid resource capture and growth, where ecological/climate conditions are (invasiveness) are opportunistic
excluding other species changing over longer time periods as conditions change e.g. tree
eucalypts or non-fire colonisers
Table 3. Ecological functional classes for CSR plant strategy schemes of Grime (2006) and equivalent Australian
species fire response strategies (Gill 1981, Keith and Tozer 2012).

8 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

tree (Casuarina cunninghamiana), a smaller root- leached of nutrients and redistributed by wind, as the
suckering, saline-tolerant tree (C. glauca), large fire- sea had gradually retreated over the previous 100 000
sensitive shrubs (Allocasuarina distyla, A. littoralis, years. The current seacoast cliffs would have been
A. verticillata) and low-growing resprouter shrubs substantial topographic features up to 150 m high,
(Allocasuarina nana, widespread; and A. diminuta and albeit 8-15 km inland (Figure 4).
A. glareicola, very localised). As a result of the size of its catchment and relative
Sites can represent local pollen catchments or a discharge, the Hawkesbury River system dominated
much wider and different habitat. Plant families with the shelf morphology and adjacent hinterland north of
abundant wind-dispersed pollen (Chenopodiaceae, Sydney. Its paleochannel formed large embayments
Asteraceae, Casuarinaceae, Poaceae, ferns and mosses) below the LGM strandline, and it strongly resembled
predominate in cores, but could have been carried a drowned river valley, relative to the outline of the
many kilometres. Aerial dispersal of material over large LGM paleo-shoreline (Albani et al. 2015).
distances can occur in dust storms and fire storms. The In contrast, from about 50 000 to 12 0000 years
chances of deposition and survival mean that many BP, Sydney Harbour would have been dry land with
species with limited pollen production and specialised a small Parramatta River paleo-watercourse forming
insect pollination are not recorded. Major groups may a narrow symmetrical gorge and sinuous path across
be missing; there are very few records of Fabaceae, the coastal sandstone shelf and draining out into
Rutaceae, Orchidaceae and Xanthorrhoeaceae, for perched freshwater swamps or seepage areas. There
example, preserved in sediments.

RESULTS
Likely vegetation on the continental
shelf coastal Plain during the Last
Glacial Maximum
The continental shelf coastal plain
About 125 000 years BP, as the milder conditions
of the Last Interglacial began to cool, sea level began
to recede, from a level about 10 m higher than today,
to a lowest level, 120 m lower than currently, at
about 20 000 years BP (Albani et al. 2015) (Figure
3). For up to 100 000 years the continental shelf off
eastern Australia, from Torres Strait to Tasmania,
was exposed to some degree, before re-inundation by
about 7 000 years ago. From about 50 000 to 12 000
years BP, the bulk of the exposed landscape is likely
to have remained a stable geomorphological feature
contributing, at its most extensive, a substantial area
of essentially terrestrial habitat, an area of about 250
000 - 300 000 ha between the Hunter and Shoalhaven
Rivers at about 18 000 years BP.
The current continental shelf visible in the
satellite images approximates the LGM coastal
plain, which extended about 8-15 km beyond the
present Sydney shoreline. The late Quaternary
paleogeomorphology from Broken Bay to Botany
Bay is described by Albani et al. (2015). The land
sloped eastward from the present coastline down
to a rocky sandstone coast, up to 20 m above the
LGM sea level. Rock platforms and outcrops about Figure 4. The coastline off Sydney between the
60-80 m in elevation (Hawkesbury and Narrabeen Hawkesbury and the Georges Rivers at the height of the
sandstones and shales) extended up to 8 km from the LGM based on bedrock geomorphology and showing
current coastline (the limit of scan surveys - Albani paleo sea level and drainage features that were active at
et al. 2015) interspersed with sandsheets and dunes, the time. (from Albani et al. 2015)

Proc. Linn. Soc. N.S.W., 146, 2024 9

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

were similar paleo-watercourses for the Botany and this time is likely to have included seagrasses and
Georges systems, and smaller drainages perched on saltmarsh, but not the characteristic mangroves of
the coastal shelf. Small sandy beaches were associated today. Mangroves are essentially tropical (Rogers et
coastal features here. al. 2017) but sea surface temperatures in the Southern
Beyond Sydney major catchment paleodrainage Ocean were 3-6 °C colder (Mooney et al. 2017), i.e.
systems included the Hunter and Shoalhaven, while about 14-17 °C for LGM Sydney, conditions likely to
smaller catchments now draining into coastal lakes have been too cold for the current Sydney mangroves,
such as Tuggerah and Illawarra are likely to have Avicennia marina (current southern limit Wilsons
drained into freshwater swamps. Present-day reefs Promontory, Victoria, is about 15-19 °C today)
and islands (e.g., Five Islands, Illawarra), would and Aegiceras corniculatum (southern limit Tuross
have added higher sandstone hills to the mostly sand Head, 17-21 °C). Mangroves are inferred to have
or sandstone landscape though some shale bedrock colonised the Sydney coastline from the north as sea
exposures are likely off the Central Coast and temperatures warmed later in the Holocene.
Illawarra. Seagrasses are readily dispersed and likely to
have had some LGM representation. Current species
Estuarine saltmarsh on the LGM coast
include temperate species of Posidonia, Ruppia
At the peak of the LGM the drowned river valley
and Zostera (Kuo et al. in Sainty et al. 2012) likely
of the Hawkesbury was about about 3 km wide and
favoured by LGM conditions. Those present at
extended for about 15 km eastwards from present-
any particular time would have depended on both
day Broken Bay (Albani et al. 2015) (Figure 4). As a
temperature conditions and local habitats.
tidal estuary it was probably fringed or perhaps filled
East coast saltmarsh flora is ecologically temperate
with sandy or silty sediments, its large catchment
and southern in origin (Adam 1990; Boon et al. 2015).
making up for the generally low LGM sedimentation
Of 53 saltmarsh dominants in Victoria, only about 25
rates (Mooney et al. 2021). Estuarine vegetation at

Figure 5. Saltmarsh species Wilsonia backhousei is mainly southern with a couple of disjunct limited occurrences as
far north as Sydney that could be remnants of a wider LGM distribution. (AVH 2022)

10 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

species also occur in NSW (G. Carr in Sainty et al. 2012), and increasingly dry microhabitats; more stress-
and of the seven main Ecological Vegetation Classes in tolerant coastal headland forms of coastal species
Victorian saltmarshes, only four occur in NSW. Unlike may have developed during this time (e.g. Actinotus
the NSW saltmarsh flora, current Victorian saltmarsh helianthi, Acacia suaveolens - Morrison and Rupp
includes shrubby chenopods and many annuals (about 1995). Though the relatively favourable coastal
38 species). During the LGM a floristically richer rainfall and temperature conditions would have
saltmarsh than today is likely for the Hawkesbury moved progressively eastward with newly exposed
estuary probably including many Victorian species; coastline, the widespread cool dry LGM conditions
disjunct northern outliers are interpreted as evidence of would have favoured now-common southern
former extent rather than recent colonisation (Table 4, shrub species (e.g. Melaleuca ericifolia (Figure 6),
Figure 5). For the smaller coastal Parramatta, Botany Melaleuca armillaris, Myoporum insulare, Acacia
and Georges River paleo-watercourses that debouched sophorae subsp. sophorae) as well as some now rare
above sea level during the LGM, saltmarsh vegetation is species such as Melaleuca biconvexa (Figure 7).
only likely to have established in the Holocene when sea These are likely to have fringed the LGM saltmarshes
level rose high enough to drown their valleys, by which and littoral habitats. The large coastal swamp
time cold–adapted southern species are likely to have forest species of today, Melaleuca quinquenervia,
retreated south. Melaleuca nodosa, Eucalyptus robusta, Casuarina
Coastal scrub and swamp forest glauca, Leptospermum laevigatum, Leptospermum
As sea level retreated with the LGM, nearby lanigerum and Leucopogon parviflorus have
existing vegetation would have colonised local bare substantially northern distributions and are likely

Figure 7. Disjunct occurrences of coastal shrub Melaleuca

Figure 6. A more northern distribution along the LGM coastal biconvexa (Port Macquarie to Jervis Bay) may be remnants of
plain is likely for the now southern coastal shrub Melaleuca a more widespread former occurrence on the alluvial flats of
ericifolia. (AVH 2022) the LGM estuarine coastal plain. (AVH 2022)

Proc. Linn. Soc. N.S.W., 146, 2024 11

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Saltmarsh species Main occurrence Disjunct northern outliers

Snapper Island (N-limit), Botany Bay, Five
Atriplex cinerea, shrub Southern Australia
Islands Illawarra
Austrostipa stipoides, grass Vict., SA., Tas., saltmarsh, seacliffs Shellharbour (N-limit), Jervis Bay
Gahnia filum, graminoid Vict., SA., common Georges River, Jervis Bay, Moruya saltmarsh
Gosford (N-limit); inland occurrences e.g. Lake
Selliera radicans, perennial herb Southern saltmarsh
Bathurst, could indicate waterbird dispersal
Tecticornia pergranulata Coastal saltmarsh, Vict. (Sainty et
Isolated up-river saltmarsh in Homebush Bay
subsp. pergranulata, shrub al. 2012).
Wilsonia backhousei,
Victoria, common Parramatta River, Jervis Bay
spreading perennial
Wilsonia rotundifolia, low Royal NP, Jervis Bay, coastal disjunct occurrences
Essentially southern Australia
spreading perennial (also Lake Bathurst and Deniliquin)

Table 4. Disjunct northern outliers of southern saltmarsh interpreted as evidence of former LGM saltmarsh
distributions.

to have been restricted to the North Coast and south onto the exposed continental shelf, and colonised
Queensland, together with their associated fauna. dune sands as they were gradually leached to extreme
Luly et al. (2010) used global climate models to infertility. At Myall Lakes on leached low-nutrient
suggest Pteropus poliocephalus (Grey headed Pleistocene sands with podsols up to 125 000 years
flying fox), and its rainforest and eucalypt LGM old, Myerscough and Carolin (1986, 2014) recognised
habitat (assuming current habitat trees, Melaleuca 13 sand communities including North Coast Banksia
quinquenervia, Eucalyptus robusta, Eucalyptus wallum, Dry heath and Dry Heath Forest. They
tereticornis and Eucalyptus siderophloia), was only describe stress-tolerator/resprouter OCBIL ecologies
a third of its modern extent, with its core distribution with lignotuber-like mass roots for Banksia aemula,
focused on the North Coast region of NSW and the and Melaleuca nodosa, and very long-lived mallee
bioclimatic envelope shifted eastwards onto the then- growth forms of Corymbia gummifera and Eucalyptus
existing LGM coastal plain. acmenoides (compared with tree forms in other sites).
Many other species were short-lived seeders. Dune-
Wallum vegetation on the LGM Coastal plain sands
swale vegetation relates to drainage and waterlogging
Remnants of the LGM coastal plain survive as
conditions supporting Wet Heath, including northern
southeast Queensland’s sand islands. Pollen cores
species Banksia oblongifolia, Banksia robur, and
from North Stradbroke Island, now dominated by
Baloskion pallens, and essentially southern species
characteristic wallum sand vegetation, indicate
Gymnoschoenus sphaerocephalus, Xanthorrhoea
floristic similarity during the LGM, with swamp
resinosa and Sprengelia incarnata in swamps.
vegetation suggesting coastal rainfall was relatively
Boundaries between vegetation on highly leached and
high at the time (Tibby et al. 2017). Further south,
younger sands are very sharp (Myerscough 2020).
from a sandy Newcastle swamp Williams (2005)
On the younger Holocene sands (< 6500 year-old)
described a sharp increase in Casuarinaceae pollen
are poorly-developed podsols, with Dry Sclerophyll
at the height of the LGM (which could be any of
Forest of Eucalyptus pilularis, Angophora costata
Casuarina glauca, Allocasuarina littoralis, A.
and Banksia serrata with Competitor/ facultative-
torulosa, A. distyla or A. verticillata), and essentially
resprouter ecologies allowing quick colonisation and
sclerophyll taxa pollen throughout the LGM period
movement onto disturbed sites.
consistent with the presence of wallum (including
Similar wallum vegetation on leached podsols,
Cyperaceae, Haloragaceae, Myriophyllum, Triglochin
fringed by young coastal dunes, is likely to have
and Restionaceae pollen relevant to the immediate
developed across the LGM coastal plain from
swamp site, and Angophora /Corymbia/ Eucalyptus
Myall Lakes to Botany Bay. At Tomago, near the
(4 types), Dodonaea triquetra, Amperea, Monotoca,
Hunter River estuary, local endemic Eucalyptus
Calytrix, Banksia, Pomaderris, Plantago, Fabaceae,
parramattensis subsp. decadens occurs on near sea
Myrsinaceae, Rutaceae, Asteraceae, Poaceae and
level Pleistocene sands with characteristic wallum
ground ferns, relevant to the surrounding area.
Banksia aemula (Bell 2022), and on similar soils
It appears likely that in the 100 000 years as sea
in endangered Kurri Kurri sand-swamp woodland
level receded following the last Interglacial, wallum
near Cessnock, which suggests a former larger LGM
species from the north would have gradually moved
coastal plain population.

12 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

South of the Hunter, highly leached Pleistocene aemula coastal wallum vegetation, now at its southern
sand is now restricted to coastal headland remnants at limits in the Eastern Suburbs (Table 5) (remnants now
Redhead, Munmorah, Bouddi National Park (Siddiqi protected as Eastern Suburbs Banksia Scrub), may
et al. 1973), North Head (80 m elevation), Long Bay have intruded farther inland, along the dry incised
and La Perouse (Eastern Suburbs Banksia Scrub). Cooks River and lower Georges River to Milperra. A
There is an inland southern occurrence at Agnes cluster of wallum species (including Banksia aemula,
Banks (25 m elevation) (Benson 1981) supporting key Baloskion pallens, Trachymene incisa, Hypolaena
wallum heath Banksia aemula (Figure 8) and other fastigiata, Ricinocarpos pinifolius, and Dillwynia
wallum species with similar northern distributions glaberrima (Figure 10)), now found on isolated
and southern limits, e.g. Baloskion pallens (Figure leached Pliocene–Pleistocene sand dunes at Agnes
9) (Table 5). This suggests southward movement of Banks (elevation 25 m) near Richmond, is consistent
vegetation such as that at Myall Lakes along a Sydney with past colonisation from the coast, particularly
LGM sandplain. during periods of strong winds and sand movement.
Sand masses off Sydney, beyond the sandstone Dune vegetation at Agnes Banks (rainfall 800 mm p.a.)
clifflines, were completely lost as Holocene sea -level has 47% of species in common with Myall Lakes and
rose, but prior sand accumulation taking place across 25% with North Stradbroke Island (Benson 1981).
the Botany Bay basin (a Rose Bay sand core dates to c. Occurrences of various coastal sand species in the
47,000 years BP - Stephen Gale, University of Sydney, Mellong-Howes Valley area may possibly be remnants
pers. comm.) would have developed a landscape of more extensive LGM dry sand vegetation.
of old leached and relatively younger sands with Coastal swamp vegetation
various wallum vegetation species, as well as exposed Evidence of wallum-associated swamp vegetation
sandstone rock platforms and gully habitats with dune, on North Stradbroke Island throughout the LGM
swale, swamp and woodland, with floristics enriched suggests coastal rainfall at the time was relatively
by southward movement. During the LGM Banksia high (Tibby et al. 2017), perhaps influenced by East

Figure 8. Banksia aemula, a characteristic wallum species reaches its southern limit on Pleistocene sand at Botany
Bay and is likely to have been important species on the LGM coastal plain. (AVH 2020)

Proc. Linn. Soc. N.S.W., 146, 2024 13

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Coast Lows (Ellerton et al. 2017). Assuming similar Gymnoschoenus in almost continuous site occupation
rainfall conditions, it is likely that similar LGM sedge for the past 22,000 years, together with Lepidosperma,
and shrub swamp vegetation occurred on the coastal Restionaceae (probably Empodisma minus),
plateaus and adjacent coastal plain farther south. Asteraceae and Chionogentias species (possibly
Pollen analyses from Machaerina/Gymnoschoenus Gentiana wingecarribiensis). Similar vegetation on
sphaerocephalus (Figure 11) moorland at 400 m poorly-drained swales of coastal Sydney sand plains
elevation on the Woronora Plateau (at Gallaghers (Eastern Suburbs) and headwater depressions on
Swamp, Avon River) (Macphail, in Hope 2010)) shows sandstone plateaus (Ku-ring-gai and Royal National

Wallum dune species Southern limit

Macarthuria neocambrica Munmorah
Acacia baueri subsp. baueri La Perouse
Bauera capitata La Perouse
Sprengelia sprengelioides La Perouse
Styphelia viridis subsp. viridis Jibbon
Banksia aemula La Perouse, Agnes Banks
Baloskion pallens Agnes Banks
Acacia quadrilateralis Eastern Sydney with an outlier at Ulladulla
Table 5. Northern Wallum ecosystem species with southern limits in the Sydney area

Figure 9. Restiad, Baloskion pallens has its southern limit on isolated hinterland Pleistocene sanddunes at Agnes
Banks, west of Blacktown, suggesting a previous more extensive occurrence on the LGM coastal plain. (AVH 2022)

14 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Figure 10. Extensive, essentially southern, coastal Figure 11. Distribution of Buttongrass, Gymnoschoenus
distribution of Dillwynia glaberrima suggests extensive sphaerocephalus highlights its essentially southern
occupation of sandy deposits on the LGM coastal plain, distribution and, north of Myall Lakes, its restriction to
but little subsequent fragmentation with sea level rise. higher (cold) habitats. Past connections along the LGM
An occurrence on sand inland at Agnes Banks, disjunct coastal plain as far as Myall Lakes are likely. (AVH
from other current coastal Sydney sites suggests LGM 2023)
connections. (AVH 2022)

Parks) today includes both widespread northern species conditions are adequate for periodic recruitment,
such as Banksia oblongifolia and Banksia robur as though they can survive with infrequent or low fire
well as essentially southern species Gymnoschoenus regimes (assuming fire events < adult plant ages).
sphaerocephalus and Sprengelia incarnata. Fire events would have been relatively infrequent
during the LGM (Mooney et al. 2011) with low
Coastal sandstone and sand habitats - Woody
temperatures, low atmospheric CO2, slow growth
obligate seeders
and little buildup of fuel, except on the wetter coastal
Woody short-lived obligate-seeders (c. 5-50
fringe where localised Aboriginal burning would have
year life spans) are ruderal /fire sensitive shrub
assisted seed recruitment episodes. Coastal conditions
species that are important components of current
may therefore have provided LGM habitats for many
sand and sandstone floras, and make up a substantial
ruderal/fire sensitive species which later spread in
component of both species and biomass in sclerophyll
warmer conditions. For example in South Australia,
heath vegetation (Keith 2004). Many have long-lived
genetic data suggest refugial populations of the
dormant soil seedbanks (many species in families
obligate seeder shrub Goodenia ovata survived on
Fabaceae, Rutaceae, Ericaceae, Sterculiaceae)
three southern peninsulas (Fleurieu, Yorke, Eyre) and
while others are serotinous and carry canopy-stored
off-shore Kangaroo Island, which formed a continuous
seed banks (Casuarinaceae and many Proteaceae,
landmass during the LGM, and later spread north
particularly the big woody shrubs such as Banksia
(Kireta et al. 2019). Goodenia ovata could have
ericifolia and Hakea teretifolia, which rely on quick
survived similarly on Sydney’s LGM coastline.
growth and abundant mass seedling recruitment
For Sydney endemic obligate-seeder shrub
for their competitiveness). Both strategies are now
Acacia linifolia, species-wide provenance boundaries
associated with episodes of mass recruitment and
(Rossetto et al. 2022) suggest recent population spread,
rapid seedling growth in good conditions (e.g. post-
probably from limited LGM coastal occurrences.
drought or post-fire) and are abundant where rainfall

Proc. Linn. Soc. N.S.W., 146, 2024 15

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Provenance studies of more widespread coastal (Briggs 1962). Subsequent Holocene warming may
obligate-seeder shrubs Acacia longifolia and Acacia have allowed some spread but their current restricted
terminalis showed large provenance boundaries (i.e. occurrences (many are endangered taxa) are likely
not localised), some level of latitudinal structuring, to have resulted from loss of favoured habitat by
and a northern boundary at the Hunter River (Rossetto competition from taller shrubs and trees, loss of LGM
et al. 2022), a pattern consistent with LGM coastal coastal plain habitat (e.g. Darwinia leptantha) and
occupation and subsequent successful spread. more frequent fire events.
In contrast, in the case of Acacia ulicifolia,
Coastal Clay soil headland species as LGM
narrower provenance boundaries with some degree of
remnants
clonality and selfing (Rossetto et al. 2022) is consistent
Shale bedrock and volcanic dykes (i.e. non-
with persistence as a stress-tolerator/resprouter
sandstone geology) exposed on the LGM coastal plain
during the LGM and little subsequent expansion.
would have supported clay soil species, and Grassy
Acacia suaveolens also showed narrow provenance
coastal headland heath (Keith 2004) is likely to include
boundaries and localised genetic clustering (Rossetto
remnants of previously more extensive LGM clay
et al. 2022) suggesting LGM persistence rather than
soil vegetation, particularly where continued coastal
recent spread. These findings align with a number
exposure has prevented later tree invasion. Stress-
of genetically fixed forms described as Acacia
tolerator Themeda triandra is likely to be a longterm
suaveolens subspecies by Morrison and Rupp (1995):
survivor. Rare low-growing ruderal shrub Pultenaea
the successful widespread modern coastal form
maritima appears to have been restricted to isolated
(subsp. suaveolens); a Myall Lakes region form with
coastal occurrences north of Terrigal by post-LGM sea
narrower phyllodes restricted to the Holocene sand
level rise, and is just surviving through a combination
dunes, and likely to have been more important on the
of open conditions and fire (Hunter 2016).
LGM coastal plain; a prostrate windswept coastal
In 1905 Linnean Society member R. H. Cambage
headlands form, that perhaps tracked the retreating
found isolated coastal occurrences of the small seeder
coastline from headlands on the LGM coastal plain or
tree Allocasuarina verticillata, a species otherwise
perhaps evolving in the rapid disturbances as coastal
from drier inland southern Australia (including Bass
headlands retreated following the post LGM sea level
Strait islands with LGM mainland connections), on
rise; while a montane resprouter form above 300 m
Narrabeen shales at Mona Vale and Stanwell Park
with longer and thicker seeds, and a small rootstock
(Figures 12 and 13). In the first publication to mention
form restricted to the Grampians region of Victoria
vegetation on the LGM coastal shelf, Cambage (1905)
are both possible LGM remnants.
concluded …having in view the isolated occurrence
Coastal sandstone as an OCBIL landscape has a
of Casuarina stricta [now Allocasuarina verticillata]
higher species richness than coastal sand, as a result
along the east coast of NSW, the suggestion that
of its longer stable history (Table 6). Darwinia taxa
it formed part of the latest flora on the present
are a group of now very restricted woody obligate-
continental shelf before its final submergence, seems
seeders needing recurring rainfall and fire events
to me less open to destructive criticism than any other
to recruit, which may have occurred more widely
put forward. Drier inland occurrences in the Hunter
in LGM sandstone coastal heath and coastal plain
Valley (Warkworth), and along the Wollondilly
vegetation. All 13 NSW taxa occur in the Sydney area
(Wanganderry, and Upper Burragorang) suggest
or South Coast (8 are endemic), all occur in heath on
coastal occurrences originated from past eastward
sandstone or shaly ironstone OCBIL habitat, though
migration in dry conditions associated with the LGM.
some have more specific edaphic requirements
Vegetation associated with the
(Briggs 1962). Some show evidence of inbreeding
sandstone plateaus and upper Blue
Mountains: evidence of in situ LGM
Taxon Myall Lakes Royal NP survival
Darwinia 1 3 Marked differences in regional climate due
Grevillea 0 11 to orographic and rainshadow effects would have
Pultenaea 6 13 influenced LGM climates. Orographic uplift is likely
Styphelia 1 5 to have extended the higher coastal rainfall zone
Rutaceae 9 25 inland onto the coastal sandstone plateaus (now Ku-
ring-gai and Royal NPs) which, though 8-15 km
Table 6. Occurrence of taxa with many seeder species at inland, with lower LGM sea level were effectively up
Myall Lakes (essentially sand habitats) and Royal NP
to 250-300 m high (Figure 14).
(essentially sandstone habitats)

16 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Figure 12. Distribution of Allocasuarina verticillata in southern Australia Figure 13. Richard Cambage recorded the
showing coastal and inland occurrences; past LGM inland and coastal shrub Allocasuarina verticillata at Newport in
plain connections are likely. (AVH 2022) 1905 and suggested it was “part of the latest flora
on the present continental shelf before its final
submergence.” (Linnean Society of NSW)

lower Blue Mountains to Yengo and northwest

across Wollemi to the Goulburn River is likely.
Average LGM temperatures are likely to have
been about 11°C (perhaps equivalent to today’s
Katoomba/Newnes Plateau conditions) (Table
7), but with more frosts. Summer insolation
and seasonality was relatively low, and low
rates of sediment accumulation confirm low
rainfall conditions (Mooney et al. 2021).
Orographic effects would have increased
rainfall in the upper Blue Mountains and
Tablelands. Montane areas would have been
colder but with less evaporation. Mean winter
temperature on the Southern Tablelands was 8
°C lower than present (Barrows et al. 2022).
Similar low temperatures are likely on the
higher Blue Mountains sandstone plateaus
above about 900-1000 m elevation and above
Figure 14. LGM areas Schematic designation of Sydney area
800 m on the Southern Highlands (Katoomba
LGM landscape regions showing likely extent across the. 1=
Coastal sandplain wallum/sandstone heath; 2=Dry mid-elevation rainfall 1091 mm pa; mean annual temperature
hinterland sandstone shrubland; 3=Subalpine sandstone heath/ 12 °C today could have had an equivalent
mallee/cliff seepage/gully complex; 4=Tablelands subalpine LGM mean temperature of 5 °C including
grassland/mallee; 5= Hunter Valley dry claysoil shrubland elevation-related adiabatic cooling). This
corridor; 6=Western Sydney grassland/shrubland. would be somewhat equivalent to Charlotte
Pass (Table 7) and a mean winter temperature
Further west, rainshadow effects are likely
of 1-2 °C. South-eastern Australia alpine areas
to have resulted in a much drier hinterland. A low
today have mean annual temperatures below 8
rainfall zone on the mid elevation sandstone plateaus
°C (Venn et al. 2017).
from the Nattai north across western Sydney, the

Proc. Linn. Soc. N.S.W., 146, 2024 17

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

COASTAL towns (elevation m) mean annual LGM estimate LGM temp. equivalent areas today
temp. °C (6 °C lower)
Port Macquarie (20 m) 18 12 Katoomba/Bombala
Newcastle 33 m 18 12 Katoomba/Bombala
Narrara 20 m 17 11 Newnes Plateau/Oberon
Sydney Obs H 39 m 17 11 Newnes Plateau/Oberon
Parramatta 55 m 18 12 Katoomba/Bombala
Campbelltown 75 m 17 11 Newnes Plateau/Oberon
Richmond 20 m 20 14 Goulburn
Valley Heights 320 m 17 11 Newnes Plateau/Oberon
Wollongong 30 m 17 11 Newnes Plateau/Oberon
Bega 50 m 15 9 Nimmitabel
Phillip Island Victoria 14.5 9 Nimmitabel
MONTANE 1 °C lower to account for LGM elevational adiabatic cooling
Newnes Plateau (1050 m) 11 4 <Charlottes Pass
Lithgow 950 m 12 5 Charlottes Pass
Katoomba 1017 m 12 5 Charlottes Pass
Bowral 662 m 13 6 Perisher
Bathurst 713 m 13 6 Perisher
Oberon 1088 m 11 4 <Charlottes Pass
Goulburn 670 m 14 7 Perisher
ALPINE
Bombala705 m 12 5 Charlottes Pass
Nalbaugh SF 675 m 11 4 <Charlottes Pass
Charlottes Pass 1755 m 5 -2 No current equivalent
Perisher 1735 m 6 -1 No current equivalent
WESTERN PLAINS
Dubbo (260 m) 17 11 Newnes Plateau/Oberon
Brewarrina 20 14 Goulburn
NORTHERN HEMISPHERE
(Britain)
London (rainfall 621 mm) 11 5 LGM - Graminoid & Forb Tundra
Edinburgh (rainfall 706 mm) 8.5 2.5 LGM Ice sheet
Table 7. An indication of Last Glacial Maximum temperatures. Current mean annual temperatures for some coastal
and montane NSW towns and estimated LGM temperatures (6-7 °C lower) and equivalent areas. Comparative
areas are not necessarily equivalent in terms of rainfall. Due to lower LGM sea-levels all sites were about 20 km
further inland and about 120 m higher.

Recent studies on scree slopes and frost cracking Stress-tolerators in sclerophyll heath - Evidence
indicate periglacial conditions during the LGM are of in situ LGM survival
likely to have affected much of southeastern Australia, Species with strong persistence traits (Bond &
down to 680 m on the Southern Tablelands (including Midgely 2001) are likely to have been favoured in
Wombeyan Caves) (Barrows et al. 2022). Enhanced the harsh LGM conditions - low stature, longevity,
effective precipitation from reduced evaporation clonal spread, drought/fire tolerance and strong
and snow melt is likely to have increased vegetation habitat fidelity (localised propagule dispersal). These
survival in montane conditions. Modelling of characteristics are also associated with species with
precipitation into snow and rain indicates that LGM a long history of in situ occupation (long before the
snowfall was a common occurrence in July across LGM) on (OCBILs) in Western Australia (Hopper
the Great Dividing Range (the Macquarie River 2021).
catchments draining much of the Central Tablelands, The stable, erosion-resistant plateaus and ridges
developed snowpack depths equivalent to 94% of of the low nutrient Sydney sandstone are similar to
present-day mean annual runoff depths) (Mueller et OCBILs, and there are many species with OCBIL site
al. undated). characteristics, essentially stress-tolerant resprouters

18 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Figure 16. Lambertia formosa distribution suggests wider

occurrence (North Coast outlier via LGM coastal plain), while
Sydney populations south of Wollongong show morphological
difference (consistently longer leaves) (PlantNet 2023). (AVH 2022)

Figure 15. Allocasuarina nana distribution (Northern limit Upper

Cudgegong River) showing extensive montane and more limited
coastal distribution, suggesting former wider LGM occurrence,
now reduced. (AVH 2019)

in sclerophyll heath and woodland vegetation cover), which is an overall distribution consistent
(Keith et al. 2004, Keith and Tozer 2012). Such with being previously more extensive in colder, drier
species include long-lived resprouter shrubs, mallee LGM conditions. Examples of other stress-tolerator/
eucalypts, prostrate/clonal groundcover species, OCBIL heath species with fragmented montane and
and rhizomatous geophytes (Keith and Tozer 2012). coastal distributions indicative of marked contractions
Many are rare and endemic species confined to heath include Allocasuarina diminuta subsp. mimica,
habitats, and likely to have occupied these sites Acacia baueri subsp. aspera, Monotoca ledifolia and
throughout the LGM. Others are more widespread. Darwinia fascicularis subsp. oligantha (Table 9).
For example, Allocasuarina nana has massive Other stress-tolerator/OCBIL shrubs are
multi-stemmed rootstocks, sheds its seeds only a widespread in both heath and woodland today,
few metres from its canopy and is abundant today including Lambertia formosa, Lomatia silaifolia,
in montane sandstone heath (Figure 15). It also has Hakea laevipes, Isopogon anemonifolius, Isopogon
limited occurrences on Sydney coastal plateaus (but anethifolius, Banksia spinulosa, Acacia ulicifolia,
only as limited patches in high light sites lacking tree Platysace linearifolia, Xanthorrhoea species and

Proc. Linn. Soc. N.S.W., 146, 2024 19

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Particular LGM Ecology atributes Advantaged growth-forms Likely life forms

conditions
Low temperature, Cold tolerance Long-lived slow-growing STRESS TOLERATORS
Low rainfall, short or multistemmed trees (resprouters) mallee eucalypts
Drought-tolerance
Low atmospheric co2,
Wind tolerance Small perennial low-growing STRESS TOLERATORS
Short growing season
or prostrate shrubs (resprouters) OCBIL shrubs,
Longevity geophytes, sedges
High wind Wind pollination, wind Mass seed release RUDERALS
dispersal Asteraceae, Poaceae
Periodic wet periods Rainfall pulse Shortlived drought dormant/ RUDERALS herbs, monocot
Open conditions recruitment seasonal ephemeral herbs
Disadvantaged growth-forms
Low temperature, Moisture, CO2 Long-lived trees Most tree eucalypts
Low rainfall, demanding) COMPETITORS
Low atmospheric CO2 Frost, snow, high wind Quick-growing woody RUDERALS Fire dependent
sensitive (through shortlived perennials seeder species,
moisture loss Mesic species
Low atmospheric CO2 CO2 requiring C4 grasses Themeda
Low fire incidence Obligate fire, smoke Shortlived fire ephemerals Obligate seeders, Fabaceae
dormancy recruitment
Table 8. Sandstone habitat life forms likely to be either advantaged or disadvantaged during Last Glacial Maximum
conditions of lower temperature, rainfall, CO2, and strong wind particularly at high elevations

Stress Tolerator/ OCBIL heath species with fragmented montane and coastal distributions
Allocasuarina nana montane sandstone heath and on the Sydney coast
Allocasuarina diminuta subsp. mimica (Blue Mountains, Bundanoon, Orchard Hills, Heathcote. Kingsford
Acacia baueri subsp. aspera Upper Blue Mountains and on the coast near Mt Keira
Monotoca ledifolia Upper Blue Mountains, Woronora Plateau and Howes Valley
Darwinia fascicularis subsp. oligantha Upper Blue Mountains, Southern Highlands and Maroota in western Sydney
Stress Tolerator/OCBIL shrubs widespread in both heath and woodland today
Lambertia formosa Sydney populations south of Bargo River show morphological difference
(consistently longer leaves)
Lomatia silaifolia Separate southern and northern populations of Lomatia silaifolia differ
genomically
Platysace linearifolia Rhizomatous capacity
Table 9. Remnants of the LGM sandstone heath landscape: Examples of Stress Tolerator/ OCBILs) in current
sclerophyll heath and woodland vegetation

many geophytic species in families Liliaceae, Mallee eucalypts – longevity, growth form and
Phormiaceae, Iridaceae, and Orchidaceae (Table 9). distributions
These have evidently been able to cope with later About 15% of the almost 100 eucalypt species
changes particularly the likely increase in tree cover in the Greater Blue Mountains World Heritage Area
competition associated with Holocene warming and (GBMWHA) are mallees (Hager and Benson 2010).
increased atmospheric CO2. Evidence of previously All are characteristic Stress-tolerator/ OCBIL species,
wider distributions becoming isolated may show up low-growing, multi-stemmed plants, with fire-
in local morphological variation. Sydney populations resistant survival and habitat fidelity, and restricted
of Lambertia formosa south of Bargo River show seed dispersal and recruitment, compared to most
local morphological difference (consistently longer tree-form species (Booth 2017).
leaves - PlantNET 2023), while the North Coast Mallees are very long-lived; more so than trees
outlier suggests wider past connections via the former in the same habitat, which are physically limited
LGM coastal plain distribution (Figure 16). Separate by an ever-growing, but increasingly scarred trunk,
southern and northern populations of Lomatia vulnerable to wind, fungal decay, termites, etc.
silaifolia differ genomically (Milner et al. 2012). One individual of the Western Australian mallee

20 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Genus/subgenus Section/series Mallee species Occurrence Habitat

Eucalyptus/ sect. Latoangulatae E. cryptica (cattai) Coast, dry ridges, Restricted, Dural Dry, low
Symphyomyrtus ser. Annulares (red rainfall
mahoganies)
Eucalyptus/ Sect. Pumilio (Grey E. pumila Hunter Valley, one site, Pokolbin, Dry, low
Symphyomyrtus gums) sandstone shrubland rainfall
Eucalyptus/ sect. Exsertaria ser. E. dealbata subsp. Hunter Valley, Dry ridges, scrubby Dry, low
Symphyomyrtus Exsertae (eastern red aperticola heath, restricted near Denman rainfall
gums)
Eucalyptus/ sect. Maidenaria ser. E. recurva Southern Tablelands restricted Dry, low
Symphyomyrtus Recurvae Braidwood rainfall
Eucalyptus/ sect. Maidenaria ser. E. pulverulenta Central Tablelands, restricted ridges Dry, low
Symphyomyrtus Viminales (white gums) Lithgow, rainfall
Eucalyptus/ ser. Pachyphloiae E camfieldii Coast, mountains heath, E. camfieldii Dry, high
Eucalyptus (stringybarks) E. bensonii and E. bensonii restricted dry ridges rainfall
E. ligustrina
Eucalyptus/ ser. Psathyroxyla E. multicaulis Blue Mtns heath, widespread Dry, high
Eucalyptus subser. Considenianae rainfall
Corymbia sect. Corymbia Red Corymbia Coast, dry heath restricted Royal NP Dry, high
Bloodwoods gummifera mallee rainfall
Eucalyptus/ sect. Maidenaria ser. E. camphora subsp. Cudgegong and Megalong Valley, Wet, high
Symphyomyrtus Foveolatae (swamp camphora (Penrose endemic), peatswamp rainfall
gums) E. aquatica margins, restricted
Eucalyptus/ sect. Maidenaria ser. E. baeuerlenii Wentworth Falls,restricted, clifflines Wet, high
Symphyomyrtus Viminales rainfall
Eucalyptus/ ser. Strictae (mallee E. stricta (sens lat.) Coast, upper BM, Sth Highlands Wet, high
Eucalyptus ashes) E. obstans / heath, periodically wet sites, mostly rainfall
burgessiana widespread
E. dendromorpha
E. cunninghamii
Eucalyptus/ ser. Pauciflorae (snow E. gregsoniana Newnes Plateau, restricted, heath Wet/high
Eucalyptus gums) periodically wet sites rainfall
Eucalyptus/ ser. Fraxinales E. luehmanniana Coast, restricted, moist sites Wet, high
Eucalyptus rainfall
Eucalyptus/ ser. Longitudinales E. stellulata Tablelands, upper BM swamp Wet, high
Eucalyptus (sallees) E. moorei periodically wet sites, frost hollows rainfall

Table 10. Mallee eucalypts in the Sydney area, occurrence and habitat from dry to wet. Classification follows (Nicolle
2021)

Eucalyptus argutifolia is estimated to be almost 2000 growth and indefinite long-term survival providing
years old (11 m across, estimated lignotuber growth habitat conditions remain stable. Mallee longevity
of 1.5–3.4 mm outwards per year based on its clonal (c. 3000-5000 years) could have allowed survival of
extent - Kennington and James 1997; Hopper 2021)). individuals through the severest peak of the LGM
Using similar measures, mallee-form individuals though the trade-offs (clonality, genomic bottlenecks)
of Corymbia gummifera in coastal heath in Royal may have reduced recovery ability (Table 10).
National Park with lignotubers up to 10.5 m across
Coastal sandstone mallees
(Mullette 1978) could be 1500-3000 years old, and
With low temperatures and low CO2, but better
southern NSW Mongarlowe mallee, Eucalyptus
local rainfall, LGM coastal vegetation is likely
recurva (with only 6 remaining individuals at 3 sites)
to have included both obligate seeder and stress-
with lignotubers up to 12 m across (Commonwealth
tolerator/resprouter shrubs including mallees. Many
Dept of Environment 2015) could possibly be 1700-
Sydney mallees are regional endemics with strong
4000 years old. Clonality has been reported in the rare
habitat fidelity; most are conspicuously associated
mallee Eucalyptus cryptica (previously called ‘sp.
with wet habitats and/or high rainfall (>1200 mm pa),
Cattai’ -Wilson et al. 2023), clonal samples in some
including the Green ash mallees (Strictae, the biggest
cases being located as much as 30 m apart (Rutherford
mallee group), snow gums and sallies, and Tableland
et al. 2022), suggesting extensive rhizomatous
swamp gums, and often associated with wet heaths

Proc. Linn. Soc. N.S.W., 146, 2024 21

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Figure 17. Mallee species A. Eucalyptus burgessiana, B. Eucalyptus dendromorpha, C. Eucalyptus camfieldii and D.
Eucalyptus luehmanniana all show disjunct occurrences indicative of former wider distributions including likely
LGM coastal connections. (AVH 2022)

(Table 10). Their current scattered distributions south) is unknown, but like northern populations
suggest previously more extensive occurrences, now of Eucalyptus burgessiana, it is still maintaining
contracted and fragmented (Figure 17). genetic connectivity despite fragmentation of habitat,
Relatively recent fragmentation events may be presumably via a range of bird and insect pollinators.
implied by small genomic differences not associated
Extensive LGM moisture-dependent mallee
with clear morphological variation. For example,
vegetation on the upper Blue Mountains
for the green ash mallee Eucalyptus burgessiana
On the basis of Northern Tablelands pollen
(including E. obstans), Rutherford et al. (2018) found
records Ellerton et al. (2017) suggested LGM montane
that though local genomic connectivity is maintained
conditions with a more positive water balance were
in populations with up to 40 km distance separations
effectively wetter than previously thought. Little
(Beacon Hill to Royal NP), population-level
Llangothlin Lagoon (1360 m) held permanent water
differentiation is evident between Sydney and Jervis
during the late LGM), and with temperatures 9-11°C
Bay populations, about 100 km apart. This is consistent
below present, supported subalpine herbfield and
with post-LGM sea level rise loss of intervening
eucalypt mallee woodland (presumably the current
LGM coastal plain populations. Populations of
frost-tolerant mallees Eucalyptus pauciflora and
Eucalyptus camfieldii and Eucalyptus luehmanniana
Eucalyptus stellulata). Llangothlin Catchment is on
in Ku-ring-gai and Royal NPs could have also been
soils derived from basalts and granites.
similarly linked through the coastal plain. However,
Applying recent climate findings (Ellerton et al.
the endemic coastal sandstone shrub Angophora
2017; Barrows et al. 2022; Mueller et al. undated) to
hispida, occupying similar habitat to these mallees
(localised low elevation patches of sandstone heath the Blue Mountains sandstone landscapes, an LGM
between Yengo and Royal NPs shows an absence of subalpine zone (>800-900 m) with a moisture-related
such population genetic structure, indicating there subalpine vegetation can be defined extending along
were no major biogeographical barriers restricting the eastern edge of the Central Tablelands from the
gene flow (Rutherford et al. 2020). Whether it ever upper Cudgegong/Coricudgy in the north, south
extended beyond its current distribution (i.e. farther through the upper Blue Mountains to Kanangra,

22 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Eucalyptus stricta Eucalyptus apiculata and E. laophila

Figure 18. LEFT: Eucalyptus stricta has a widespread distribution associated with heath and rocky sandstone OCBIL
landscapes and like Allocasuarina nana, with a relatively narrow coastal band, consistent with previous more extensive
occurrences in LGM conditions. RIGHT: The narrower leaves (phenotypic plasticity) and drier sites for Eucalyptus
laophila (Newnes Plateau, Wollemi National Park), and Eucalyptus apiculata (west of Wollongong) (both in E. stricta
genomic envelope) at current elevational limits may indicate some post-LGM montane elevation expansion. (AVH 2022)

and east along the Southern Highlands to Robertson mallees are subject to only brief snowfalls but the
(Figure 14 - Area 3). This area is surprisingly mallee habit with small crowns on tall very flexible
consistent with the distributions of the moisture- stems would appear resilient to heavier winter
dependent sandstone mallees, stringybark mallees snowpack
on better-drained soils (Table 10), and most of the Subalpine vegetation on the Upper Blue
stress-tolerator/resprouter shrub species such as Mountains sandstone plateaus and cliffs
Allocasuarina nana and Lambertia formosa, as well At higher elevations, a surprisingly diverse
as smaller prostrate montane heath species, many LGM landscape is likely as a result of local climatic
of which appear to be indicative of previously more conditions and geomorphology. Bare ground or
extensive occurrences across this area. open vegetation on dry exposed sandstone at high
Eucalyptus stricta, for example, the most elevations with high wind velocities is suggested
widespread of the Sydney mallees, is associated with from remnant sand-dunes on the Newnes Plateau
rocky sandstone heath and wet sites near peat swamps active during the LGM (Hesse et al. 2003); perhaps
and, like Allocasuarina nana, with relatively few drier conditions in the northwest of the region at this
coastal occurrences and more extensive population time isolated the higher areas of Coricudgy and the
concentrations on the upper Blue Mountains, Southern upper Cudgegong.
Highlands and Budawang area (Figure 18), consistent The high daisy pollen abundance recorded from
with widespread past landscape predominance. LGM Tablelands pollen cores is not reflected in any
Mallees with similar sandstone habitats but more Asteraceae richness in current sandstone heath. Most
limited distributions are Eucalyptus gregsoniana montane shrubs today are in stable sites with reliable
and Eucalyptus moorei (Table 10). Today montane periodic moisture, with features of benefit in LGM

Proc. Linn. Soc. N.S.W., 146, 2024 23

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Alpine species Lifeform Habitat Substrate Fertility Locality

Asplenium flabellifolium fern moist ss low BM, CSS

Empodisma minus graminoid wet ss low BM, NP, CSS

Gonocarpus micranthus herb wet ss low BM, NP, CSS

Asperula gunnii herb wet ss low BM, NP
Celmisia longifolia herb wet ss low NP
Celmisia tomentella herb wet ss low BM, KAN
Baeckea utilis shrub wet ss low KAN, NP
Epacris paludosa shrub wet ss low KAN, NP
Epilobium gunnianum herb wet ss low KAN, NP
Blechnum pennamarina fern wet ss low KAN
Myriophyllum pedunculatum subsp.
herb wet ss low NP
pedunculatum
Epacris microphylla shrub wet-dry ss low BM, NP, CSS
Stylidium graminifolium herb wet-dry ss low BM, NP, CSS
Oxylobium ellipticum shrub dry ss low BM
Carex gaudichaudiana graminoid wet shale med BM, NP
Neopaxia australasica herb wet shale basalt med KAN
Rhodanthe anthemoides herb dry shale ss med BM, CP, CSS
Viola betonicifolia herb dry shale med BM, CP
Crassula sieberiana herb dry shale ss med BM, CP, CSS
Microseris lanceolata herb dry shale basalt med KAN
Pimelea ligustrina shrub dry shale basalt med BM, KAN
Geranium potentilloides herb dry shale basalt med BM,
Lagenifera stipitata herb dry shale basalt med-high BM, NP, CP
Carex breviculmis graminoid dry shale basalt med-high BM, CP
Melicytus dentatus [Hymenanthera] shrub dry shale basalt med high BM, KAN
Oreomyrrhis eriopoda herb dry shale basalt med high BM, NP,
Polystichum proliferum fern dry shale basalt high BM
Figure 11. Distribution of Buttongrass, Gymnoschoenus sphaerocephalus highlights its essentially southern
distribution and, north of Myall Lakes, its restriction to higher (cold) habitats. Past connections along the LGM
coastal plain as far as Myall Lakes are likely. (AVH 2023)

conditions such as low spreading or prostrate growth Isopogon prostratus has a series of localised small
forms, slow growth and longevity features such as disjunct populations on sandstone landscapes from
root-suckering, frost and snow tolerance, temperature- Newnes Plateau along the Southern ranges to Nadgee,
induced flowering and short growing-season responses. and a final southern occurrence in Victoria (Benson and
Now restricted species such as Phyllota squarrosa, von Richter 2010) (Figure 19). Its seed, dispersed from
Mirbelia platyloboides, Persoonia hindii, Isopogon serotinous fruits in the leaf litter, is limited to within a
prostratus and Kunzea cambagei are likely to have been metre or so of its cones, maintaining long-term habitat
more abundant on the treeless LGM uplands. Many fidelity but with very little colonising potential. In open
have patchy or fragmented populations in the upper windy LGM conditions, they may have moved larger
Blue Mountains and/or Southern Highlands, and farther distances (African Proteaceae seeds can move up to 500
south. For example the prostrate sclerophyll shrub m (Bond 1988).

24 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

relative, the similarly poorly-dispersing, partly

clonal Pherosphaera hookeriana, a major LGM
vegetation component in Tasmanian. The latter now
persists in small populations directly descended from
nearby glacial populations (Macphail, et al. 2014;
Worth et al. 2021).
Now-restricted Blue Mountain angiosperm
species associated with moist sandstone cliff-line
habitats include stress-tolerator OCBIL shrubs
Eucalyptus baeuerlenii, Isopogon fletcheri, Epacris
hamiltonii and Epacris apiculata; geophytes Alania
cunninghamii, Blandfordia cunninghamii (Coleby
Figure 19. The prostrate resprouter shrub Isopogon 2006), and rhizomatous herb Euphrasia bowdeniae.
prostratus is likely to have been important in the open Species in periodically waterlogged habitats include
LGM heath. Its groundlevel flowers and serotinous Boronia deanei, Dillwynia stipulifera and Olearia
fruits keep its populations very localised, but its quercifolia, the clonal Haloragodendron gibsonii
sporadic occurrences from Newnes Plateau to Victoria (Sydes and Peakall 1998), ruderal/fire-sensitive
suggest a former more continuous LGM distribution. Sprengelia monticola, Leionema lachneoides, and
(Photo Ian Baird) Acacia ptychoclada as well as more common species
including ferns (Gleichenia rupestris) and monocots.
Links with the Kosciuszko alpine flora are few. These all have requirements for permanent moisture
Of about 200 current Kosciuszko alpine (i.e. above but could have survived in local mesic LGM
the treeline) taxa (Costin et al. 1979), only about 14% microhabitats.
(27 species) also occur in the Sydney area (Table 11). Though current groundwater-dependant montane
About half of these occur on relatively dry fertile peat swamps only date from about 12 000-10 000
habitats on the Great Divide consistent with the LGM years BP, with similar dates on the Woronora Plateau
shrub or grassland steppe vegetation of the pollen (Young 2017), the big water-dependent but stress-
studies (discussed below). The other half, about 14 tolerator tussock sedges occurring on the LGM coast,
species, occur in wet bog or swamp habitats in the Gymnoschoenus sphaerocephalus, Xyris ustulata
Blue Mountains, many on Newnes Plateau (Table and Lepidosperma limicola may have also survived
11). Many of these are separated by long distances the drier LGM subalpine conditions confined to
from their main Kosciuszko occurrences (e.g. locally moist seepages with snow melt and enhanced
Celmisia longifolia and Celmisia tomentella). Other effective precipitation from reduced evaporation
taxa have similar but less fragmentary disjunctions, (Barrows et al. 2022),. There is no evidence that more
some reaching as far as the Northern Tablelands. extensive Gymnoschoenus Buttongrass moorland (as
In the LGM subalpine landscape, water inception in Tasmania) was part of the LGM subalpine Blue
rather than surface run-off is likely to have been Mountains vegetation, nor whether any particular
important (Mueller et al. undated). Moisture run-off species were predominant.
into aquifers of alternating sandstone and shale strata Were species at high elevations limited by lack of
(predominating features of the sandstone geology) pollinators?
would have emerged as cliff-line seepages and Evidence here is very limited. It is likely that
springs, providing habitat for small alpine shrubs, most plant species’ survival in high elevation LGM
herbs and ferns. conditions was primarily limited by the harsh
The lack of Kosciuszko connections and the growing conditions. Windy conditions would have
survival of its own wet cliff-face and groundwater- assisted wind-pollinated plants such as members of
dependant assemblages is consistent with a separate the Casuarinaceae, Asteraceae and Poaceae.
long vegetation history for the Blue Mountains. The limited literature on LGM climate and bird
A classic example is the palaeoendemic dioecious distributions relate to continental scales or rainfall
conifer Pherosphaera fitzgeraldii, now restricted to (e.g. Magpie-(Toon et al. 2007); Shrikethrush (Lamb
a few waterfall-spray habitats near Katoomba (Smith et al. 2019); butcherbirds (Kearns et al. 2014)), rather
1981), with plants showing a very limited genetic than regional elevation responses that may impact
variation across its range (Wyn Jones pers. comm.). pollination. Honeyeaters are important pollinators for
It may have been more widespread during the LGM, many species of Proteaceae, Fabaceae and Myrtaceae;
and contracted subsequently, like its only surviving current autumn migration routes take them along

Proc. Linn. Soc. N.S.W., 146, 2024 25

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

the coast and into the upper Blue Mountains up to being impacted by hybrid swamping (Rutherford et
at least 1000 m elevation. Such routes are likely to al. 2022).
depend on flower source availability (Munro et al. Eucalyptus dealbata subsp. aperticola is
1993). Assuming LGM flowering was in the warmest possibly a remnant of an LGM dry country invasion
seasons, plants could have been sought out and by Eucalyptus dealbata sens. lat. (subsp. aperticola
pollinated during bird migration. Shrub-pollinating occurs within 40 km of subsp. dealbata). The
mammals including bush rats and possums are dry LGM conditions would have enabled drier
likely to have survived in LGM heath. Mountain western species to invade Sydney sandstone.
Pygmy Possum, Burramys parvus, now confined Dry country shrubs Calytrix tetragona, Gaudium
to alpine habitats at Mt Hotham and Kosciuszko as parvifolium, Acacia triptera, Leucopogon muticus,
well as the Blue Mountains is likely to have been Boronia anethifolia, Isopogon dawsonii, Harmogia
separated earlier than the LGM, and has a need for densifolia, Cryptandra spinescens, Gompholobium
rocky boulder habitat with low temperatures. This aspalathoides and Allocasuarina gymnanthera were
would have rendered most of the intervening region recorded with Eucalyptus dealbata subsp. aperticola
unsuitable, even in times of glacial maxima (Hoser (Bell and Nicolle 2020). All are low-nutrient shrub
2020). Insects, particularly native bees (Halictidae, species (many are obligate seeders) with essentially
Colletidae), are important for Ericaceae and Fabaceae north-western NSW occurrences, e.g. Narrabri
(Johnson and McQuillan 2011; Bernhardt et al. and Pilliga. Some species, such as Allocasuarina
2019). While there were probably enough pollinators luehmannii, appear to have spread eastward as far as
for generalist plant species to survive, species the lower elevation sandstone of the Goulburn River
dependent on particular insect interactions (e.g. some and Wollemi/Yengo National Parks, some to the
terrestrial orchids) may have been restricted to lower coastal Sydney sandstone plateaus (e.g. Leucopogon
elevations, due to limitations of temperature and wind muticus) and presumably along the LGM coastal
on specific insect survival. Some now rare remnant plain south to Nowra (e.g. Eucalyptus beyeriana
LGM stress-tolerator shrubs are self-fertile (e.g. (Figure 20), Gaudium parvifolium, Cassinia
Haloragodendron gibsonii - Sydes and Peakall 1998) cunninghamii). Phylogenomic studies of Calytrix
and able to maintain seed production in absence of tetragona identified almost 20 infraspecific groups
outcrossing opportunities or pollinators. across southern Australia, suggesting vicariance and
long-term site fidelity (Nge et al. 2021). Research on
Dry sandstone country: species invade from the west
some other common, widespread, and often variable
On the mid-elevation essentially dry rainshadow
species may disclose similar patterns of variation.
sandstone areas between the coast and the higher
Obligate seeders are not likely to have been
Tablelands (Figure 14 - Area 2), without the
advantaged in LGM conditions, particularly away
elevational benefit of snowpack and reduced lower
from the coast where warmer wetter conditions
evaporation, the cold dry windy LGM conditions and
and some degree of Aboriginal burning is likely.
lower atmospheric CO2, could probably only support
However, in drier hinterland areas with little fuel
dry scrub or shrubland. In this area, mallees are rare
build-up and rare fires, long stable periods could have
(Table 10) though small trees of Angophora bakeri
helped some larger woody obligate seeders with a
and Corymbia eximia are LGM possibilities. Of the
combination of ruderal and drought stress-tolerator
few mallee species occurring there now, Eucalyptus
traits. Common Western Slopes and Tablelands
pumila and Eucalyptus dealbata subsp. aperticola
conifer Callitris endlicheri has extensive western
are restricted to scrubby heath at a few sites on the
Blue Mountains occurrences and a few isolated
southern rim of the Hunter Valley (Bell and Nicolle
eastern populations (Coxs River, Burragorang
2020). There is no suggestion of previously wider
Valley and Darkes Forest - a coastal sandstone heath
occurrences such as the Green Ashes, though in
Endangered population (NSW Scientific Committee
the case of Eucalyptus cryptica, now restricted to
2004)). Callitris rhomboidea and Callitris muelleri
sandstone ridges at Dural-Annangrove, Rutherford
have widely separated upper Blue Mountains
et al. (2022) found varied genetic diversity across
and coastal (Mosman, Chatswood-Terrey Hills)
populations and unexpectedly high clonality via its
distributions, as well as disjunct occurrences in
underground lignotuber. This is consistent with a past
the Southern Highlands (Callitris muelleri) and at
more extensive local population, reduced, perhaps in
Nowra (Callitris rhomboidea). These occurrences
the early Holocene, by more vigorous invading red
are consistent with more extensive distributions in
mahogany trees, and then persisting through clonal
LGM and post-LGM warming conditions but, as fire-
longevity on isolated low nutrient ridges. It is now
sensitive seeders, it is likely ENSO-related increased

26 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

E. racemosa and E. globoidea.

Unlike the montane mallees they
all currently occupy well-drained
sites, are quick growing with
rapid seedling recruitment and are
unlikely to have benefitted in low
LGM CO2, winter snowpack and
waterlogged soils. All could have
possibly survived harsh LGM
conditions, at least in sheltered
sites; nearby prior site occupation
and rapid expansion with post-
LGM warming may account for
their predominance as trees at
higher elevations today.
Eucalypt pollen identified by
Robbie and Martin (2007) from
about 19 000 years BP at Mountain
Lagoon (500 m elevation) in the
lower Blue Mountains includes
sandstone habitat Eucalyptus
piperita and E. racemosa, shale
country E. crebra, or perhaps
sandstone habitat E. beyeriana
though specific identification
of eucalypts even with living
material may be difficult. Tree/
mallee Angophora and Corymbia
species, Angophora costata,
A. bakeri (except Rutherford’s
Kings Tableland A. bakeri mallee)
Figure 20. Distribution of Eucalyptus beyeriana suggests eastward spread via the and Corymbia gummifera have
lower elevation sandstone Goulburn River and Wollemi/Yengo NPs and coastal northern distributions, and appear
sandstone plateaus, with the LGM coastal plain enabling spread south to Nowra. more temperature-sensitive based
(AVH 2022) on their generally low-elevation
restricted ranges (<900m). They
fire/drought conditions over the last 6000 years have are thus more likely to have been confined to the
confined occurrences to the current localised rocky LGM coast. The likely spread of dryland species
ridges, harbour headlands or riparian sites that act as with clay soil affinities, e.g. Eucalyptus sideroxylon,
fire refugia. Martin (1994) reports Callitris pollen at E. fibrosa, E. moluccana and E. crebra, into western
Kurnell from 8000-5000 years BP, but reduced levels Sydney at the same time along nearby river systems
thereafter. is discussed below.
Mallee/tree species and Sheltered Eucalypt Persistence of rainforest and mesic vegetation
Woodland concentrations Rainforest species richness is highest at tropical
Harsh conditions or frequent fire today may and subtropical latitudes. Subtropical and northern
reduce some essentially tree-form eucalypts to stunted Warm Temperate assemblages have predominantly
trees, though they rarely form true multi-stemmed Queensland and northern NSW distributions (e.g.
mallees. Most are in Eucalyptus subgenus Eucalyptus Gondwana Rainforests World Heritage Area) but
with essentially southern temperate distributions (i.e. extend with decreasing floristic richness as far south
northern limits coinciding with Sydney sandstone as the Shoalhaven River (Metcalfe and Green 2017).
limits) –including Eucalyptus sieberi, E. piperita, Cool Temperate assemblages occur in southern
E. consideniana, E. racemosa and E. scias. Some NSW, Victoria, and Tasmania. Rainforest habitat
have very strong drought resilience (Zolfaghar et al. is generally restricted to higher nutrient soils than
2015) for example Eucalyptus sieberi, E. piperita, soils derived only from sandstone: on basalt caps

Proc. Linn. Soc. N.S.W., 146, 2024 27

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

in the Blue Mountains, some shale outcrops in the Refugial limitations may have restricted
Permian sandstones or exposed in gullies in the subsequent expansion for some species. Genetic
Blue Mountains and along the coast, volcanics in the evidence indicates that during the LGM and earlier
Illawarra, and some locally enriched coastal alluvial glacial periods, Atherosperma moschatum in south-
soils (Keith 2004). eastern Australia was reduced to four separate
During the LGM suitable east coast habitat regions, including the northern Barrington Tops/
for many rainforest species including Doryphora Upper Blue Mountains, and Monga on the South
sassafras (Figure 21), Elaeocarpus reticulatus, Coast; followed by local dispersal in the Holocene
Callicoma serratifolia, Tristaniopsis collina, (Worth et al. 2011). Cool Temperate Nothofagus
Tristaniopsis laurina and Pittosporum multiflorum cunninghamii occurred in the northern Kosciuszko
was less in extent, on average 65%, than occupied at region 35 000 years ago as indicated by old wood
present (Das et al. 2019). In the Sydney area, sheltered (Caine and Jennings 1968). The species is now
Blue Mountains gullies or coastal sites on fertile soils restricted to Victoria and Tasmania where current
would have provided LGM refugia for these mesic populations are derived from short-range dispersal
species as well as cool adapted Temperate rainforest (about 100 km) from nearby LGM glacial refugia
species including Ceratopetalum apetalum (Heslewood (Worth et al. 2009).
et al. 2014). Multiple stems from swollen stem bases Conditions on the LGM coast would have been
in woody species such as Atherosperma moschatum, milder than in the mountains. Eucryphia moorei
Eucryphia moorei and Doryphora sassafras (Johnstone (Figure 22) is likely to have survived the LGM
and Lacey 1983) would have provided persistence in coastal sites such as the Illawara and extended
and extended longevity in LGM conditions. Other into Warm Temperate rainforest later. The recent
Temperate tree species can occur as multi-stemmed discovery of the local endemic and long-lived
shrubs (e.g. Ceratopetalum apetalum) suggesting that clonal tree species Daphnandra johnsonii (and an
LGM mesic refugial pocket vegetation may have been endemic gall midge apparently dependent on its
as scrub rather than as rainforest. seed) on volcanics in the Kiama area (Lemmon
2019; Kolesik et al. 2019) suggests long-term site
persistence before, and presumably throughout, the
LGM on higher nutrient, old, climatically-buffered
relatively fertile OCFEL landscapes (Hopper et al.
2021). This is a parallel situation to sclerophyll
OCBILs). Disjunct occurrences of some dry
rainforest climbers (stress-tolerator resprouters with
extensive root spreads, and possibly clonal), on
clay soils in coastal Wollongong and inland western
Sydney (Razorback, Cobbitty) and Hunter Valley
sites (such as Cynanchum elegans at Mt Dangar,
Razorback, Shellharbour and Marsdenia viridiflora
at Razorback) may be remnants of LGM dry scrub
vegetation. These are more genetically diverse in
southern areas than Indo-Malesian lineages (such
as the recent Indo-Malesian migrant Toona ciliata)
with a recent history of north to south migration
(Rossetto et al. 2018) (Figure 21).
Tough, vegetatively spreading but moisture-
sensitive ferns such as Calochlaena dubia, some
Blechnum, Microsorum and Asplenium species.
are likely to have survived the LGM with reduced
distributions. In Europe, Asplenium polyploid
complexes can indicate glacial refugia (Vogel et al.
1999). LGM refugia have been shown to be important
Figure 21. Habitat suitability models for LGM, Mid for South Island New Zealand ferns (Marske and
Holocene and current periods for cool adapted Doryphora Boyer 2024). The larger long-lived trunked ferns
sassafras (A) contrast with a recent Indo-Malesian migrant, Dicksonia antarctica, Cyathea australis (considered
Toona australis (B) (adapted from Rossetto et al. 2018). hardy in cultivations to -6 °C) and Todea barbara

28 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Rainforest (Mills 2023), with similarities to

the North Coast Nightcap-Border Ranges
and Dorrigo areas identified as paleo-refugial
areas of high genetic diversity (Rossetto et
al. 2018), and presumably OCFELs, based
on studies on rainforest species in Royal
National Park and Barrington Tops, were
identified as mostly recolonised areas by
Rossetto et al. (2018).

The Central Tablelands and Great

Dividing Range
Treeless alpine daisy-rich grassland and
mallee woodland
On the Tablelands mean winter
temperature during the LGM was 8 °C colder
than present with subalpine conditions down
to 680 m (Barrows et al. 2022). The Great
Dividing Range area (Oberon, Bathurst,
Bowral, Goulburn, at 650-1100 m elevation)
is likely to have experienced, cold (4-7 °C
subalpine/alpine conditions (Table 7). South-
eastern Australian alpine areas today have
mean annual temperatures below 8 °C (Venn
et al. 2017).
In contrast to the sandstone Blue
Mountains, much of the Tablelands is
metamorphic in geology with higher nutrient
clay soils than the sedimentary Sydney Basin.
Pollen analyses for similar clay landscapes
in south-eastern Australia (Lake George and
southern Victoria) indicate LGM vegetation
of Asteraceae, Chenopodiaceae, Myrtaceae
and Poaceae, and has been interpreted
variously as xerophytic shrub, heath and
Figure 22. Southern cool temperate tree Eucryphia moorei is likely woodland, or semi-arid grassland steppe
to have survived the LGM in coastal sites such as the Illawarra (Pickett et al. 2004; Mooney et al. 2017).
and extended into warm temperate rainforest subsequently. Is Chenopodiaceae is indicative of drier inland
its disjunct record at Springwood west of Sydney a remnant or a winter rainfall areas often on calcareous soils
chance migrant? (AVH 2022) or perhaps local saline areas, and with low
occurrence on the Tablelands today may
have wide distributions in creeks and moist forest represent largely wind or dust-storm transported
in the Blue Mountains and along the Tablelands and pollen. However, during the LGM, grassy/low shrub
coast from Queensland to Tasmania but isolated vegetation with many current Tablelands perennial
remnant inland populations including Mt Kaputar grasses and herbs is likely to have occurred across
(near Narrabri) for Dicksonia, Rockley (south the drier Tablelands, with much of the Bathurst Plains
of Bathurst) for Cyathea, and Mingham Springs essentially treeless in the 19th century (Semple 1997).
(west of Orange) for Todea suggest the drier LGM This applies also to the grassy clay soil woodlands
conditions reduced previous wider distributions. of western Sydney, where Poaceae and Asteraceae
Increased Holocene rainfall may have expanded are still well-represented. Fossil Asteraceae pollen
Blue Mountains populations. The Illawarra-Southern appears related to Cassinia (Chalson and Martin
Highlands is a particular refugial area with 51% 2008), relatively short-lived coloniser shrubs in the
of NSW fern species occuring in the Minnamurra area today. Of about 200 current Kosciuszko alpine

Proc. Linn. Soc. N.S.W., 146, 2024 29

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

(i.e. above the treeline) taxa (Costin et al. 1979) only Tops, Coolah Tops and New England farther north,
about 14% (27) also occur in the Sydney area and coincides with areas of likely LGM subalpine to
about half of these occur on relatively dry fertile alpine conditions. The present Eucalyptus stellulata
habitats on the Great Divide (Table 11), consistent ecology and moist frost-hollow habitat suggest its
with LGM grassy/shrub vegetation observed in occurrence as part of an extensive LGM Central
pollen studies. Tablelands moisture-dependent subalpine mallee
The enhanced effective precipitation resulting woodland. Occurrences of other now restricted,
from low temperatures, reduced evaporation and mostly poorly-drained habitat mallee/small
snow melt suggested for the LGM at montane trees provide support for this view (Eucalyptus
elevations on the Northern (Ellerton et al. 2017) and camphora subsp. camphora (Cudgegong and
Southern Tablelands (Barrows et al. 2022) is also Megalong Valley); Eucalyptus aggregata (Bathurst-
applicable to the Central Tablelands where similar Wallerawang, Southern Highlands); Eucalyptus
LGM moisture-dependent swamp and subalpine aquatica (Southern Highlands swamp endemic);
woodland is likely. The current distribution of and Eucalyptus macarthurii (Boyd Plateau,
small tree/mallee Eucalyptus stellulata (Figure Southern Highlands)). Other LGM mallee woodland
23) (frost hardy to -14 °C) across the Tablelands, species probably included Eucalyptus dives current
together with isolated occurrences on Barrington distribution of which probably aligns well with the
extent of LGM alpine woodland on non-
sandstone landscapes across south-east
Australia) and Eucalyptus pauciflora
(widespread, cold-tolerant to -23 °C,
but low drought tolerance). Highland
populations of Eucalyptus pauciflora
in Tasmania appear to have originated
from lowland refugia rather than from in
situ high-altitude refugia (Gauli et al.
2014).
Some of the Kosciuszko alpine
species in the Sydney area are bog or
swamp habitat species, consistent with
the LGM swampy, gravel floodplain
habitat, surrounded by alpine daisy-
rich grassland, reported for Southern
Tablelands Micalong Swamp at 980 m
elevation (Kemp and Hope 2014).
Other species likely to be remnants
of an LGM mosaic of moist subalpine
mallee woodland and montane daisy-
rich grassland include cold-tolerant
shrubs such as Grevillea acanthifolia
and Grevillea juniperina, with adult
growth tolerating temperatures of
-12 °C reported in northwest USA
horticulture). Both of these species
have a series of allopatric essentially
montane subspecies, suggesting a widely
distributed taxon in LGM conditions that
has subsequently fragmented (Table 12).
A further case is seen in the localised
Figure 23. Distributional extent of the small tree/mallee Eucalyptus endemics Callistemon megalongensis
stellulata approximates the area of LGM alpine grassland shrubland and Callistemon purpurascens in similar
along the Great Divide inferred from soil/geomorphology. Now wet habitat in Megalong Valley. None
confined to wet sites and frost hollows it is a suggested remnant of of these species gives the impression of
such landscape conditions during the LGM. (AVH 2022) benefitting in the current climate.

30 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Grevillea taxa Distribution Growthform Habitat

Grevillea acanthifolia NT Tenterfield - Low spreading shrub Heath near streams
subsp. stenomera Werrikimbe
subsp. acanthifolia CT Blue Mtns shrub Peatswamps
subsp. paludosa ST Nalbaugh Shrub Sphagnum bogs
Grevillea juniperina subsp. NT-NWS Prostrate shrub
allojohnsonii
subsp. juniperina CC Western Sydney Shrub Woodland
subsp. sulphurea CT western Blue Mtns Weakly erect shrub Streamside or riparian
Coxs R-Tallong
subsp. trinervis CT Hampton-Bindo Ck Spreading low shrub ?Woodland
subsp. villosa ST Braidwood Erect shrub Riparian
subsp. amphitricha ST Braidwood- Nerriga Prostrate shrub Woodland, grassland
subsp. fortis ACT Molonglo R Robust shrub Above permanent watercourses
Table 12. The allopatric subspecies of Grevillea acanthifolia and Grevillea juniperina are mostly shrubby or prostrate
and associated with moist cold sites mainly on the Tablelands, and suggest a widely distributed taxon in LGM conditions,
subsequently fragmented. USA horticulture suggests survival in temperatures as low as -12 °C for both species.

Western Sydney landscapes and the LGM and western Sydney area, particularly in shale forests,
The fertile landscapes of the Cumberland Plain: than is evident in the present remnant vegetation.
LGM Treeless shrubland/grassland vegetation Pollen core Poaceae could include C3 perennial
During the LGM Western Sydney’s mean grasses (Dichelachne, Elymus, Microlaena, Poa)
temperature was probably about 11°C, equivalent to and C4 perennial grasses (Aristida, Bothriochloa,
today’s Upper Blue Mountains (Table 7), and in the Sorghum, Sporobolus, Themeda), all of which
rainshadow of the adjacent mountains and farther from occur today in Cumberland Plain Woodland and
the coast, much drier with rainfall of perhaps 300-400 in Tableland habitats. The herb and geophyte
mm p.a. compared with today’s 600-800 mm (Figure groundcovers in Cumberland Plain Woodland today
14 - Area 6). Such conditions would have been too include many non-woody stress-tolerator /resprouters
dry for today’s dominant Cumberland Plain trees and with persistent rootstocks allowing survival through
shrubs. Based on a 38,000 year Castlereagh pollen core drought and fire. Brunoniella australis, Eremophila
(Chalson and Martin 2008) considered this to have debilis, Plantago gaudichaudii, Asperula conferta
been a treeless LGM shrubland/grassland, similar exhibit ecologies that would have also coped with dry
to the semi-arid grassland steppe reported for the and windy conditions, frosts, ice and slow growth in
Tablelands and southeastern Australia more broadly lower atmospheric CO2, and they probably benefitted
(Kershaw et al. 1991). Most pollen identifications are from the absence of tree canopy competition.
limited to families, such as Poaceae and Cyperaceae.
Connections with drier Western NSW for clay
Chalson and Martin (2008) attributed the abundant
soil species
daisy pollen Tubuliflorites pleistocenicus to Cassinia
Drier periods would have allowed western
arcuata or Calomeria amaranthoides but this needs
species to move eastward. Isolated populations
further elucidation. Schmidt-Lebuhn and Constable
of Western tussock grass Triodia scariosa in
(2013) include both these species in a narrowly-
the Capertee Valley (Mt Gundangaroo) suggest
circumscribed clade with a range of Sydney Cassinia
eastwards spread in dry LGM conditions, and
species including Cassinia arcuata, C. aculeata,
fragmentation in the following wetter Holocene. The
C. aureonitens, C. cunninghamii, C. compacta, C.
Hunter Valley immediately north, though a barrier to
denticulata, C. laevis, C. longifolia, C. quinquefaria
north-south movement for rainforest and sclerophyll
and C. trinervia. Many of these species now occur
flora, provided a low elevation low rainfall corridor
sporadically, and are often rare, across Sydney from
between the Western Slopes clay soils and the
coast to montane habitats mainly on medium to
Newcastle- Cessnock hinterland, allowing medium-
higher nutrient soils (basalt shales or shale-enriched
nutrient-requiring western species to penetrate
sandstone) on disturbed/post-fire sites. Their likely
eastward. Bell and Driscoll (2016) list 30 western
success as colonizing species with some wind
mid layer shrubs and understorey species in the upper
dispersal under past conditions of continuous habitat
Hunter Valley, many of which extend eastwards to
may have been overlooked. The limited 19th century
the Muswellbrook and Singleton districts including
records suggest a greater abundance in the now urban

Proc. Linn. Soc. N.S.W., 146, 2024 31

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

No of species in Cumberland Plain remnant likely to be lost as annual rainfall decreases

current flora
700-800

Figure 24. About 40 % (50 species) of the native flora in a Mt Annan Cumberland Plain Woodland site also occur in
western areas with rainfall as low as 300 mm pa.

Figure 25. Movement of species on fertile shale basalt corridors (now mostly cleared) in western Sydney and the Hunter;
pre and post LGM inland connections (yellow arrows); southward LGM movement along LGM coastal plain (green);
and post-LGM coastal invasion of competitor trees and shrubs into Cumberland Plain and Hunter Valley (blue).

32 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

A B C
Figure 26. Distributions of Cumberland Plain Woodland groundlayer species A. Pimelea glauca; B. Eremophila debilis; and C.
Phyllanthus virgatus, indicate western clay-soil connections through the Southern Highlands (Mittagong, Wombeyan Caves) and
Southern and Central Tablelands (Hunter Valley), probably in more extensive grassland during the LGM or previous interglacial;
indeed, the severe LGM conditions at the high elevations may have triggered the fragmentation. (AVH 2023)

Eucalyptus camaldulensis, Allocasuarina luehmannii, layer species suggest previous, mostly now broken,
Allocasuarina verticillata. The colonising abilities, western connections. For example, Sida corrugata,
as well as prolific pollen production of many Calotis lappulacea and Glossocardia bidens have
Casuarinaceae, may explain their abundance in widespread western occurrences while Phyllanthus
pollen records. Likewise the arid-zone and western virgatus has occurrences connecting with the
Acacia species, including A. aneura, A. harpophylla, northwest. Brunoniella australis, Cymbonotus
A. melvillei, A. homalophylla, A. salicina, and A. lawsonianus, Arthropodium minus and Lotus
spectabilis. Remnant populations of some may australis have connections with Coxs River,
indicate eastern incursions isolated and outcompeted extending to Kanimbla Valley. The now rare pea
during subsequent wetter climatic periods. Swainsona monticola and small Cumberland Plain
Many Cumberland Plain groundcover species Woodland shrub Pimelea glauca occur at Bathurst.
also occur in the Temperate Montane Grasslands Zornia dyctiocarpa has disjunct populations in
(Keith 2004) and drier Tablelands and Western both the Illawarra and Southern Tablelands and
Slopes (Benson and Howell 2002). A Cumberland an extensive subtropical - tropical distribution
Plain Woodland remnant at Mt Annan has (Figure 26). None of these species occurs on the
about 50 such species (40 %) that also occur surrounding sandstone OCBIL landscapes.
in western areas with rainfall as low as 300 mm The general lack of endemism in the Cumberland
p.a. (Figure 24). Many Cumberland Plain herbs, shale groundcover flora (particularly in comparison
grasses and woody shrubs have wind- or animal- with the Castlereagh Tertiary alluvium flora)
dispersed seeds, allowing movement along habitat suggests relatively recent invasion, and though
‘corridors’ and between ‘islands’ of medium or many species have faunal-assisted seed dispersal,
high nutrient shale, basalt and alluvial soil habitats. others do not. There are some local endemics
Though isolated edaphically by the infertile Blue including Grevillea juniperina subsp. juniperina,
Mountains sandstone plateaus, western Sydney with its six related subspecies on the Tablelands
would have been, and still is, connected through (Table 12) it has a similar distribution pattern to the
shale soil habitats through the Southern Highlands groundcover species. In contrast Pimelea spicata,
(Mittagong-Robertson) and Wollondilly- a small local endemic herbaceous resprouter, has
Wombeyan Caves-Taralga areas with the Southern its main distribution across the Cumberland Plain
Tablelands and Burragorang-Coxs Valley to the and a separate isolated population at Shellharbour
Bathurst plains, areas of grassland habitat during on the Illawarra coast (Figure 27). Yapp et al.
the LGM (Figure 25). (2019) found little between-population genetic
Disjunct distributions of western Sydney ground connectivity in P. spicata with individuals within

Proc. Linn. Soc. N.S.W., 146, 2024 33

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

The low nutrient sandy clays on the ancient

Tertiary alluvials
Tertiary sediments in the Windsor-Castlereagh-
Penrith and Milperra-Liverpool-Holsworthy areas,
as well as South Maroota, have characteristic
vegetation with some localised species. Soils are
low-nutrient sandy dispersible clays, sometimes
with ironstone gravels. Broadly described as
Sydney Sand Flats Dry Sclerophyll Forest and
considered unique to the Sydney-Newcastle area
(Keith 2004), the vegetation is quite different from
the adjacent Wianamatta Shale Cumberland Plain
Woodland; there are relatively sharp boundaries
and few species in common (Benson 1992). The
western area today is the driest part of Sydney,

Figure 27. Local endemic Pimelea spicata occurs on shale

soil across the Cumberland Plain, but surprisingly the
most northern occurrences are genetically most similar to
the disjunct southern coastal Shellharbour occurrences.
Given the coastal position at Shellharbour, a possibility
is that during the LGM Pimelea spicata also extended
eastward from the Hawkesbury across the more mesic
shale areas of northern Sydney and onto the coastal plain,
with southern extension to the Illawarra. (AVH 2023)

populations similar to each other but differing

from those at other sites, findings consistent with
relatively long in situ presence across Western
Sydney. However, surprisingly, they found the
most northern occurrences (Freemans Reach and
Wilberforce were genetically most similar to
the disjunct southern Shellharbour occurrences,
and clearly differentiated from the intervening
central Cumberland Plain populations. Given
the coastal position at Shellharbour, a possibility
suggested by the genetics is that during the LGM
Pimelea spicata also extended eastward from the
Hawkesbury across the more mesic shale areas
of northern Sydney and onto the coastal plain on
outcropping Narrabeen Group & Permian strata Figure 28. Acacia pubescens occurs as scattered root
with southern extension to the Illawarra. With -suckering clusters on low nutrient clays from the lower
increased Holocene warming it may have been Blue Mountains to the Georges and Cooks Rivers in the
outcompeted by increasing Turpentine-Ironbark south, and Oakdale northwest of Wollongong, suggesting
Forest shading, and isolated at Shellharbour by survivors of previously more extensive populations. A
rising sea level. coastal outlier at South Nowra may indicate previous
LGM coastal plain connections. (AVH 2022)

34 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

Species Distrib. Habitat Grime/ fire Likely LGM habitat

ecology
Acacia pubescens west Syd. disj. Low nutrient clay ST/Res LGM open heath or open shrubland
Allocasuarina glareicola Castlereagh Low nutrient clay ST/Res LGM open heath or open shrubland
Eucalyptus sideroxylon west Syd. disj. Low nutrient clay ST/Res LGM western invading tree
Micromyrtus minutiflora Castlereagh area Low nutrient clay Rud/ ?Kss LGM desert ephemeral
Dillwynia tenuifolia Castlereagh, Low nutrient clay Rud/ Kss LGM desert ephemeral
Maroota, Howes
Valley
Pultenaea parviflora Castlereagh area Low nutrient clay Rud /Kss LGM desert ephemeral

Prostanthera scutellarioides Castlereagh area Low nutrient clay Rud /Kss LGM desert ephemeral

Grevillea juniperina subsp. west Syd broad pop Low nutrient clay Rud- Comp/ LGM shrubland
juniperina Kss

Dodonaea falcata Castlereagh, Bondi, Low nutrient clay Rud/ ?K LGM desert ephemeral
Howes V. disj.

Darwinia biflora Maroota shale /ss ironstone Rud/ Kss LGM ephemeral
-Turramurra

Table 13. Rare or regionally restricted species on Low Nutrient Clay habitats in western Sydney, showing possible
LGM occurrence/ecology. All are likely to have declined with warmer wetter conditions and increased competition.
Grime categories ST=Stress Tolerant, R=Ruderal; Fire resp. Res= resprouts, Kss=killed, seeder; Regional pop.
disj=disjunct pops.

in the rain-shadow of the Blue Mountains and species include local endemics Dillwynia
coastal plateaus, with the dryness probably tenuifolia, Pultenaea parviflora, Micromyrtus
accentuated during the LGM by lower rainfall and minutiflora, Persoonia nutans and Prostanthera
increased distance from the coast. The essentially scutellarioides, mostly species with limited seed
sclerophyll flora includes endemics: a mixture of distance-dispersal (consistent with long term
stress-tolerator/resprouter shrubs and geophytes, OCBIL site occupation). Some have disturbance-
consistent with an OCBIL landscape (its elevation related ecologies involving rainfall events for
of 15-40 m was above the 10 m sea level height at successful recruitment such as Micromyrtus
the beginning of the last Interglacial). It also has minutiflora, and are able to establish from soil
ruderal/fire-sensitive shrubs, and trees and shrubs seedbanks without fire (though heat and smoke
with wider western connections. significantly increase germination) but need open
The stress-tolerant resprouter shrubs include conditions to do so (Bangel et al. 2023). Survival
restricted local Castlereagh endemic shrubs in LGM conditions, especially for obligate-seeder
Allocasuarina glareicola and Acacia pubescens Fabaceae species, could have involved very long
(Figure 28) and more regionally widespread Acacia seed dormancy periods, alternating with shorter
bynoeana, Macrozamia spiralis and Xanthorrhoea periods as adults, effectively a desert-ephemeral
minor. All are likely to have coped with the ecology with reduced need for regular rainfall.
dry LGM conditions and can be interpreted as For Dillwynia tenuifolia at least, fruiting success
survivors of previously more extensive LGM open from both cross- and self-pollination events
heath or open shrubland (Table 13). Most now (Rymer et al. 2002) would allow build-up of soil
occur sporadically in small populations but have seedbanks, allowing rapid exploitation of disturbed
strong soil habitat-fidelity for low-nutrient sandy areas. In the drier LGM conditions, with low CO2
to sandy dispersible clay soils, or somewhat shaly and reduced biomass, a very low fire frequency,
ironstone cappings; these populations are likely to perhaps a once per century event might have been
have benefitted from LGM conditions. enough to reset populations. On the basis of its
The ruderal/fire sensitive obligate-seeder population genetics Rymer et al. (2002) suggested

Proc. Linn. Soc. N.S.W., 146, 2024 35

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Figure 29. Dillwynia tenuifolia

is now restricted to particular
low nutrient clay habitats but
population genetics suggests
the species was previously more
continuously distributed across its
range. (AVH 2023)

Figure 30. Eucalyptus sideroxylon

occurs in drier Western Slopes
woodland, in particular Gilgandra
(near Dubbo) and Pilliga
Scrub (north of Dubbo) (areas
with substrate similarities to
Castlereagh) and may have moved
east into western Sydney in drier
LGM or early Holocene conditions.
(AVH 2022)

36 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

that Dillwynia tenuifolia may have formerly been perhaps eastward invasion associated with the
more continuously distributed over its current LGM, when its current drought-sensitivity (Payne et
range (western Sydney, South Maroota, Howes al. 2012) may have been countered by lower LGM
Valley), and subsequently retreated to its current precipitation/evaporation conditions.
distribution of separate populations (Figure 29). In contrast to the other Castlereagh woodland
If so, an overall population linking disparate sites trees, Angophora bakeri is also common on low
(including Kemps Creek, Castlereagh, Marsden rainfall Dharug-Yengo, lower Blue Mountains and
Park, South Maroota, and Howes Valley) across western Woronora sandstone as well as the sandy
the intervening sandstone country, is likely during clay sites. Occurrence during the LGM included
the LGM or early Holocene climatic conditions. montane areas - a genomically distinct Angophora
Subsequent retreat on all but low nutrient clay bakeri mallee population survives in upper Blue
sites, outcompeted by tree canopy and tall shrub Mountains heath (Rutherford et al. 2021).
growth in warmer wetter conditions is a possibility. In summary it is possible that a pre-LGM low
Different histories are likely for the Castlereagh open sclerophyll shrub vegetation at Castlereagh
tree species. Eastward invasion in drier LGM or included stress-tolerant species with OCBIL origins,
early Holocene conditions may have allowed as well as obligate-seeder ruderals with more
Eucalyptus sideroxylon, E. fibrosa and E. crebra, widespread distributions on ironstone cappings and
which now mainly occur in drier Western Slopes sandy residuals, as at Kemps Creek, Liverpool-
woodlands such as Gilgandra and the Pilliga Scrub Milperra-Birrong areas, Maroota and Mellong –
(areas with substrate similarities to Castlereagh), Howes Valley. Invasion by drier western tree and
to invade western Sydney (Figure 30). Eucalyptus shrub species associated with dry LGM conditions
crebra/beyeriana pollen is reported from Mountain is likely. All are likely to have declined subsequently
Lagoon from about 19 000 years BP (Robbie and with warmer wetter conditions and increased
Martin 2007). These trees and occurrences of competition from coastal species.
small, also western shrubs (e.g.
Melaleuca diosmatifolia and
M. thymifolia) at Castlereagh
and in the Coxs-Burragorang-
Wollondilly Valleys, Capertee
Valley, or associated with
the Hunter Valley, indicate
availability of more or less
continuous clay soil/alluvial
habitat passages through the
sandstone landscapes. Eucalyptus
fibrosa, E. crebra, Melaleuca
diosmatifolia and Angophora
bakeri, for example, occur
together on Shoalhaven Group
shales in the Kedumba Valley
(Keith and Benson 1988).
Eucalyptus parramattensis
subsp. parramattensis occurs on
low elevation Central Western
Slopes to the coast (Figure 31) on
poorly-drained periodically wet
sites with very low nutrient sandy
or sandy clay substrates. It occurs
in the Goulburn and Capertee
River valleys with Eucalyptus
sideroxylon, E. crebra and E. Figure 31. Distribution of disjunct occurrences of Eucalyptus parramattensis
racemosa, as it does at Castlereagh, subsp. parramattensis, and subspecies decadens, near Newcastle. Now very
and suggests a previously more restricted subspecies decadens is likely to have been more extensive on the
widespread distribution, with now-lost LGM coastal plain. (AVH 2022)

Proc. Linn. Soc. N.S.W., 146, 2024 37

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

DISCUSSION Though not immediately observable, such patterns may

be seen with genetic studies, which will probably lead
This work began as a personal quest for
to a reassessment of some traditional species. Events
understanding the distribution of rare species and
leaving their imprints on geomorphological features,
whether an ecological history open to hypothesis
such as the emergence and submersion of the LGM
testing could be developed. Of necessity this work
coastal plain, are pivotal for timing events but have been
had been an exercise in conjecture supported by
rarely invoked in relation to species patterns.
circumstantial evidence and I am surprised at how
much has been able to be fitted together to provide A Sydney perspective that can be expanded
a reasonably satisfying account. This work has been approached from a
Sydney region perspective. That similar historical
Assuming the LGM as the major event
vegetation changes occurred concurrently north and
The extreme climatic conditions of 20 000 years
south of Sydney is evident in similar distribution
ago were not new to the landscape. There had been a
patterns for analogous species on similar geological
series of alternating eight glacial periods in the last
substrates. For example, botanists have long
800 000 years (Jouzel et al. 2007) with components
observed that on the Southern Tablelands, and on the
of the vegetation presumably responding to each shift
chain of montane islands extending from Sydney to
by moving up or down climatic gradients. The LGM
Gippsland (Victoria) (Table 14) including Newnes
would have exacerbated the vegetation impacts of the
Plateau/Blue Mountains; Hilltop-Mittagong-
preceding ones and has left the most impressionable
Penrose; Nerriga-Budawang; Deua-Wadbilliga; and
impact. Cycles of population expansion, fragmentation
Bombala-Nalbaugh many relatively rare shrub taxa
and isolation would have already happened for many
turn up on sites with similar low-nutrient (OCBIL)
species as a result of the local extinctions and bottlenecks
(mostly sandstone) landscapes. These sites are
of earlier glacial cycles. Throughout the work questions
generally isolated from each other by hundreds
of timing, age and rate of species change, arise.
of kilometres (Figure 32). The distance-dispersal
Morphological differences are the basis of traditional
characteristics and habitat similarity for many of
taxonomic interpretation (species, subspecies, forms
these species suggest similar histories in response
etc.) but they do not necessarily pick up more subtle
to contemporary climatic cycles as suggested for the
variation developed over periods of 10 000 years.
Sydney area.

Southern Tableland connections NP uBM Mitt Nerr Deua Bom SM Vic

STRESS TOLERATORS
Eucalyptus stricta Mallee x x x x
Eucalyptus baeuerlenii Mallee x x x
Boronia deanei - 2 subsp. Shrub x x x x
Genoplesium superbum geophyte x x
Acacia kybeanensis shrub x x x
Dillwynia stipulifera shrub x x
Isopogon prostratus Prostrate x x x x x x
Mirbelia platyloboides Prostrate x x
Kunzea cambagei Prostrate x x
Phyllota squarrosa Prostrate x x
Phyllota humifusum Prostrate x
RUDERALS
Sprengelia incarnata shrub x x x x x x
Grevillea renwickiana Prostrate x

Table 14. Examples of disjunct species with montane connections between Sydney and Victorian border.
NP =Newnes Plateau, Clarence, uBM =Upper Bl Mtns, Mitt =Hilltop, Mittagong, Penrose, Nerr =Nerriga, Mongarlow,
Budawangs, Deua =Deua National Park, Bom =Bombala/ Nalbaugh 1000 m SM =Snowy Mtns, Vic =Victoria

38 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

mesic species. The coastal sandplain would have

also provided a more continuous corridor (now
broken) for northward and southward movement of
oligotrophic (OCBIL) taxa, allowing connections
between the Sydney Basin with the Moreton Basin
sandstones around Grafton (the North Coast Dry
Sclerophyll Forests of Keith 2004), which have
a remarkable number of taxa sister to Sydney
sandstone endemics. Connections from there to
Gibraltar Range and other leucogranite outcrops
(Northern Escarpment Dry Sclerophyll Forests
and Northern Montane Heaths of Keith 2004)
where sclerophyll endemic taxa with similar
OCBIL characteristics to the Sydney sandstones
have developed (e.g., Gibraltar Range National
Park - Clarke and Myerscough 2006) suggest
similar parallel histories with the Sydney area.
For example, interesting parallels are evident
with related mallees. Gibraltar Range swamp-
endemic Eucalyptus dissita (section Eucalyptus
series Longitudinale - Nicolle 2021), is a habitat
counterpart to the closely related Blue Mountains
wet heath Eucalyptus moorei subsp. moorei
(Sheringham et al. 2021). The related Eucalyptus
stellulata also occurs there but is more widespread.
Swamp gum Eucalyptus camphora subsp. relicta
(Eucalyptus subgenus Symphyomyrtus section
Maidenaria; counterpart of southern Eucalyptus
camphora subsp. camphora) is associated with
Figure 32. Major disjunct montane sandstone areas in closed wet heaths and Sphagnum Bogs in Warra
southern NSW. 1 Newnes Plateau; 2 Blue Mountains; 3 National Park.
Southern Highlands; 4 Nowra-Nerriga; 5 Nadgee. Phylogenetic analyses of northern species of
series Strictae mallee ashes (Rutherford et al. 2015)
McDougall et al. (2023) report the isolated
placed three northern NSW species Eucalyptus
Devonian quartzite/sedimentary rock summit of Mt
approximans, E. codonocarpa and E. microcodon,
Imlay (886m elevation) near Eden is home to the
Sydney region Eucalyptus cunninghamii, and southern
endemic mallee Eucalyptus imlayensis. Two close
NSW-Victoria border species Eucalyptus kybeanensis
relatives occur nearby, Eucalyptus elaeophloia,
and E. paliformis in a single clade, though the
only on high-elevation Nunniong Plateau in East
northern, Sydney and southern species are in isolated
Gippsland, and Eucalyptus baeuerlenii on the
patches at least 300-400 km apart geographically.
sandstone island chain Deua-Wadbilliga, Nerriga-
Rutherford (2020) suggested vicariance (speciation
Budawang and upper Blue Mountains. The restricted
following population isolation) as an explanation.
distributions of these Eucalyptus section Maidenaria
These distributions indicate past vegetation histories
series Viminales mallees (Nicolle 2021) parallel the
for Southern and Northern Tablelands low-nutrient
Sydney area section Eucalyptus series Strictae Green
montane landscapes paralleling the Sydney area;
mallees and support the regional significance of
a bigger picture incorporating both needs to be
mallee eucalypts as glacial era survivors.
developed.
To the north of Sydney, substrate limitations
(the northern end of Sydney sandstone, the shale/ Remnant LGM sites and in situ conservation
alluvial soils of the Hunter River Valley and Interesting sites with concentrations of likely
Barrington volcanics) are barriers to movement of LGM refugial species are recognisable (Table 15).
the sclerophyll shrub flora but coastal connections Some are already recognised for their endemic flora,
associated with the LGM coastal plain have allowed though their conservation significance as evidence of
southward movement for (animal-dispersed) vegetation history has not been previously identified.

Proc. Linn. Soc. N.S.W., 146, 2024 39

 VEGETATION PATTERNS ACROSS THE SYDNEY BASIN

Landscapes with LGM Examples Example species Vulnerability issues

remnants
Moist sandstone Clifflines Upper Blue Mtns Fire refugia Pherosphera fitzgeraldii increased warming and fire,
OCBIL weeds
Sandstone mesas, Newnes Plateau; Genowlan Pultenaea praecipua Competition from canopy
ridgetops, mallee heath Point heath (Capertee Valley) Isopogon prostratus shading
OCBIL (NSW Scientific committee)
Ridgetops with clay Mangrove Mtn, Warrah trig, Grevillea caleyi Small, fragmented areas
residuals and/or remnant Duffys Forest, Maroota,
ironstone cappings OCBIL Menai, Heathcote, Waterfall
Widespread sandstone Long lived resprouter shrubs Allocasuarina nana, Recruitment limitations
plateau woodland OCBIL often widespread Lambertia formosa with frequent fire
Pleistocene leached sand Redhead, Munmorah Bouddi Banksia aemula Small, fragmented areas
sites Remnants of the LGM North Head la Perouse,
Coastal plain flora Agnes Banks, Kurri Kurri
sand swamp WL
Coastal Themeda grassy Low nutrient clays Themeda triandra, Small, fragmented areas
headland, OCFEL Allocasuarina verticillata
Low nutrient sandy clay Castlereagh, Windsor Micromyrtus minutiflora, Small, fragmented areas
vegetation, Tertiary Penrith, Milperra; Kurri sand Allocasuarina glareicola
alluvium, OCBIL swamp woodland
Western Sydney groundcover species, often Pimelea spicata, Brunoniella Lack of recognition, small
Cumberland Plain with vegetative spread australis geophytes size
Woodland Medium nutrient
shale OCFEL
Mallee swamp woodland, Western Bl Mtns, Penrose Eucalyptus camphora, E. Small size
OCFEL stellulata, E. aquatica
Western Bl Mtns Medium Box woodland Basalt caps Triodia scariosa, Small, fragmented areas
nutrient clays, OCFEL and diatremes groundcover species
Dry country shale and Native Vineyard Cobbitty Long-lived vines, Small, fragmented areas
volcanics with Dry Hunter valley remnants Cynanchum elegans Weed infested
rainforest and vine thicket,
corridor remnants, OCFEL
Table 15. Landscapes with likely concentrations of remnant or persistent species suggesting Last Glacial Maximum
in situ occurrence. All occur as small fragmented areas.

Most are OCBILs, sandstone or low-nutrient clay of species/site associations. The recognition and
sites, but others are parts of more fertile shale OCFEL continuing association of other species with particular
landscapes (Wianamatta and Permian Shales) with sites is important as associated taxa such as invertebrates
assemblages including long-resident faithful species and fungi may be similarly restricted and relict (Moir et
(as well as including trees and shrub invaders). Though al. 2011).
these sites do not show the extent of local endemism
Conservation responses for out of climate phase
evident on the sandstone OCBILs, probably because
species
the main families (Asteraceae, Poaceae) have much
The recognition that vegetation communities
better propagule dispersal, the groundcover flora
may include clusters of species with different histories
includes many species with extensive root suckering
has implications for conservation assessment and
or rhizomes (such as Brunoniella australis) consistent
management. Low-nutrient OCBIL landscapes that have
with very long association with sites.
remained stable and vegetated for very long periods may
The continued occurrence of rare species in their
include faithful species that have evolved in association
natural wild sites has significant scientific and heritage
with the site though past periods of major climatic
value for understanding historical landscape changes. Ex-
change. Actual species behaviour under different past
situ conservation (seedbanks, etc.) is being increasingly
climates is unknown, especially where there are no
developed for many localised or restricted species, but it
analogous climatic sites today. Such species may not
is important to recognise in-situ conservation measures
necessarily have characteristics associated with current
are necessary to retain the biogeographical significance
ecological health indicators, such as large seedsets,

40 Proc. Linn. Soc. N.S.W., 146, 2024

 D. BENSON

vigorous seedling recruitment or high genetic diversity. genomic studies interpreted in historical contexts,
They may, however, retain abilities for coping with and long-term field studies, is needed. In practice,
cold or drought beyond necessary requirements today. different species are likely to have had more complex,
Species that have coped with LGM conditions may and much longer histories than the relatively simple
retain considerable resilience, not necessarily evident story postulated here.
from their narrow remnant sites. In relatively stable Finally, this project raises a host of questions
but limited habitats, such as ridges and mountain tops, relating to other elements of the LGM biodiversity,
survival may be through attributes such as continued especially the biota intimately connected with plants,
underground vegetative expansion (essentially for example specialized insect-pollinator relations as
continuous lignotuber expansion, or localised spread with orchids, and with mycorrhizal and other fungi.
via root-suckering, e.g. Banksia paludosa (Baird and Are these all subsequent blow-ins or survivalists?
Benson 2022). These may allow species to persist Where were vertebrates and invertebrates (particularly
during periods when seedset or recruitment conditions those sensitive to temperature) during the LGM?
are unfavourable, and presumably evolve new forms
and ecologies if reproduction recommences. Many of ACKNOWLEDGEMENTS
these species such as mallees may be very long-lived,
The Australasian Virtual Herbarium is acknowledged
perhaps thousands of years. for its role in making available much of the data necessary
Successful seed production is often an important for this paper. The author also acknowledges all those
conservation management measure for Endangered colleagues who have at various times over many years
Species Recovery plans, but seed production may speculated about past vegetation distribution, the ecology
be difficult to achieve in many naturally restricted of restricted species and possible causes of rarity, and
species that may be clonal or partly clonal. Pressure many other things, particularly while out in the field.
to intrude genetically by translocation of individuals Particular thanks to David Keith for helpful and supportive
from other sites may introduce pathogens and do commentary and Susan Rutherford, whose genetic research
provided an impetus for the work. Thanks also to those
little genetically for populations of long-lived plants
people whose work I have dissected for insights and
that have survived past changes despite restricted perhaps taken beyond their considered limits. I take full
propagule production, provided these natural responsibility for interpretations made in this paper, and
populations remain undisturbed. hope that by stimulating interest in likely past scenarios,
As well as the restricted (mostly listed as rare or awareness of the significance of current vegetation and
endangered) slow-growing woody shrub resprouters species patterns and the importance of their conservation,
species, postulated as likely relics, there are others will be increased.
with widespread occurrence and longevity which
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