The Eucalypts of the Greater Blue Mountains World Heritage Area: distribution, classification and habitats of the species of Eucalyptus, Angophora and Corymbia (family Myrtaceae) recorded in its eight conservation reserves

Tim Hager1 and Doug Benson2

1Metropolitan Branch, Environment Protection and Regulation Group, Department of Environment, Climate Change and Water, Hurstville 2Botanic Gardens Trust, Royal Botanic Gardens, Mrs Macquaries Rd, Sydney 2000 AUSTRALIA

Abstract: The Greater Blue Mountains World Heritage Area (GBMWHA), immediately west of Sydney (33° 53’S; 151° 13’E), on the east coast of Australia was listed as World Heritage for its outstanding natural values, a major component of which is the high number of eucalypt species and eucalypt-dominated communities present, some 13 per cent of all eucalypt species in the world. They grow in a great variety of plant communities, from tall closed forests, through open forests and woodlands, to stunted mallee shrublands. This paper provides a definitive list of the 96 eucalypts (species of the genera Eucalyptus, Angophora and Corymbia in the family Myrtaceae), that have been recorded there (55 widespread, 41 restricted), together with the distribution of the eucalypts in the eight reserves that make up the GBMWHA, and information on the classification and habitat of the different species. The information is based on records held at the National Herbarium of New South Wales and the results of surveys by the NSW Department of Environment, Climate Change and Water (DECCW) over the past 20 years. The majority of species have components of both stress-tolerator and competitor ecological strategies and this has probably been a main contributor to their success. However details of the ecology of the majority of taxa are poorly-known and more research is needed to provide guidance for conservation management in the face of changing climate conditions. Cunninghamia (2010) 10(4): 425–444

Introduction • For containing the most important and significant natural habitats for in-situ conservation of biological The Greater Blue Mountains World Heritage Area diversity, including those containing threatened species (GBMWHA) immediately west of Sydney (33° 53’S; 151° of outstanding universal value from the point of view 13’E), on the east coast of Australia, was inscribed on the of science or conservation. World Heritage list in 2000. It covers over one million hectares It is not just because of its magnificent scenery, or the distant (1 032 649 ha) and covers eight protected conservation blue haze that sometimes appears over the rugged landscape. reserves, largely contiguous but partly separated by transport It is the eucalypt forests, more than anything else, that make and urban corridors. There are seven national parks as well the area special. The Blue Mountains have been described as as the Jenolan Caves Karst Conservation Reserve (2 422 ha). a natural laboratory for the study of evolution of eucalypts. The National Parks are Blue Mountains (247 840 ha in area), In the mountains’ diverse plant communities, you can trace Wollemi (499 879 ha), Yengo (153 483 ha), Nattai (47 855 the changing nature of the Australian environment – from ha), Kanangra-Boyd (65 379 ha), Gardens of Stone (15 150 geological shifts and climate variations, through to the ha) and Thirlmere Lakes (641 ha) (Figure 1). Approximately impact of Aboriginal settlement and European colonisation. 60% of the area is dedicated as wilderness (669 000 ha). The aim of this paper is to provide a definitive list of the The GBMWHA was listed for its outstanding natural GBMWHA eucalypts (species of the genera Eucalyptus, universal values: Angophora and Corymbia in the family Myrtaceae), together • As an outstanding example representing significant with information on their distribution, classification and habitat ongoing ecological and biological processes in the which will provide a basis for further research and to assist evolution of terrestrial ecosystems and communities of communication of the important biodiversity values of the area. plants and animals; and

Cunninghamia: a journal of plant ecology for eastern Australia © 2010 Botanic Gardens Trust www.rbgsyd.nsw.gov.au/science/Scientific_publications/cunninghamia 426 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

Eucalypts in Australia charcoal deposits (suggesting that fire was a factor even then). However, they remained a minor component of the Tertiary The eucalypts or gum trees are a characteristic Australian rainforest until about 20 million years ago when the gradual group of species (mainly trees but with some small shrublike drying of the continent and depletion of soil nutrients led to mallees) belonging in the Myrtaceae family (for ecology the development of more open forest types, predominantly of Myrtaceae in the Sydney area – see Myerscough 1998). Casuarina and Acacia species. Major redistributions of More than 98% of the 700 or so known eucalypt species eucalypt forest occurred in southeastern Australia during are endemic to Australia. Most eucalypts are in the genus the Quaternary, probably in relation to both temperature and Eucalyptus but the similarly-looking and closely-related aridity changes. According to McKinnon et al (2004), patterns genera Angophora and Corymbia are also traditionally of morphological variation within Eucalyptus in Tasmania, included as eucalypts in general usage. Eucalyptus itself in particular intergradation between species, suggest that is considered to include two large subgenera, Eucalyptus these redistributions might have enabled divergent species to (Monocalyptus) and Symphyomyrtus, and a number of come into contact and exchange genes through hybridization smaller ones (notably Eudesmia in northern Australia) (reticulate evolution). Similar processes probably occurred in As well as the bud and fruit characteristics which are generally the GBMWHA leading to closely-related groups of species needed for positive identification, a distinctive characteristic such as the Eucalyptus–Green-leaved Ashes. of the group is the nature of the bark. Bark types which give their names to broad groups include stringybarks, ironbarks, boxes, bloodwoods, peppermints as well as smooth-barked The Eucalypts of the Greater Blue Mountains gums. These bark types are the basis for many of the common World Heritage Area names applied to the species though unfortunately they do not necessarily align with the generic or subgeneric groups. A list of eucalypt species present was included in the Smooth-barked gums for example occurring in most of the GBMWHA nomination in 1998 (Appendix 6.2 in NPWS major groups e.g. Angophora costata, Corymbia maculata, 1998). This list has now been updated to include newly Eucalyptus tereticornis. described species and delete unconfirmed species, and enlarged to provide species occurrence in the individual Eucalypts originated between 35 and 50 million years reserves of the GBMWHA with data from specimen records ago, not long after Australia-New Guinea separated from held at the National Herbarium of New South Wales, together Gondwana, their rise coinciding with an increase in fossil

Fig. 1. Proportion of area of individual conservation reserves making up the Greater Blue Mountains World Heritage Area (total area one million hectares) Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 427

Table 1a. Eucalypt species with relatively widespread distributions in the Greater Blue Mountains World Heritage Area showing occurrence in conservation reserves. Based on records from the National Herbarium of New South Wales and surveys by the NSW Dept of Environment, Climate Change and Water.

National Parks Wollemi Yengo Gard. of Blue Jenolan Kanang. Thirl. Nattai Stone Mtns Karst Boyd Lakes Cons Res

Reserve size (ha) 499879 153483 15150 247840 2422 ha 65379 641 47855 Eucalypt species

Angophora bakeri W Y B T N Angophora costata W Y B N Angophora floribunda W Y G B K T N Corymbia eximia W Y B T Corymbia gummifera W Y B K T N Eucalyptus agglomerata W Y B K T N Eucalyptus albens W G B N Eucalyptus amplifolia subsp. amplifolia W Y B Eucalyptus bicostata W B J Eucalyptus blakelyi W G B Eucalyptus blaxlandii W G B J K N Eucalyptus bosistoana B K N Eucalyptus bridgesiana W B Eucalyptus consideniana W B Eucalyptus crebra W Y G B K N Eucalyptus cypellocarpa W G B J K N Eucalyptus dalrympleana subsp. dalrympleana W G B J K Eucalyptus deanei W Y B K N Eucalyptus dives W G B J K Eucalyptus elata B K N Eucalyptus eugenioides W Y G B J K N Eucalyptus fastigata W G B J K Eucalyptus fibrosa W Y G B K N Eucalyptus globoidea W B N Eucalyptus macrorhyncha W G B Eucalyptus mannifera subsp. gullickii W G B K Eucalyptus mannifera subsp. mannifera G B K Eucalyptus melliodora W Y G B J K N Eucalyptus moluccana W Y G B N Eucalyptus multicaulis W G B Eucalyptus notabilis Y B Eucalyptus obliqua B K Eucalyptus oreades W G B N Eucalyptus ovata B K N Eucalyptus paniculata subsp. paniculata W Y B Eucalyptus parramattensis subsp. parramattensis W Y G T N Eucalyptus pauciflora B J K Eucalyptus pilularis Y B Eucalyptus piperita W Y G B K T N Eucalyptus polyanthemos subsp. polyanthemos W G Eucalyptus praecox W G Eucalyptus punctata W Y G B J K T N Eucalyptus radiata subsp. radiata W G B J K N Eucalyptus resinifera subsp. resinifera W B T Eucalyptus rossii W G B Eucalyptus saligna W Y Eucalyptus sclerophylla W Y G B K N Eucalyptus sideroxylon W B K N Eucalyptus sieberi W G B J K T N Eucalyptus smithii B K Eucalyptus sparsifolia W Y G B J K N Eucalyptus stricta W B K Eucalyptus tenella W G Eucalyptus tereticornis W Y G B K N Eucalyptus viminalis W G B J K N 428 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area Cons. listing IUCN ROTAP,

2RC-, VU D2* 2RC-,

3RC 3RCa

2RCa VU D2* 2RCa 2VCi

2RCa VU D2 * 2RCa

2RCa LC *

2VC-VU D2 * 2RCa Cons. listing- statutory

Vulnerable (TSC) Vulnerable

Vulnerable (TSC) Vulnerable (EPBC) Vulnerable

Vulnerable (TSC) Vulnerable (EPBC) Vulnerable Vulnerable (TSC) Vulnerable

Act; *=IUCN (proposed by Bell 2008). Notes 2008). Bell by (proposed *=IUCN Act; EPBC Act, EPBC=Commonwealth Act, TSC Conservation listings TSC=NSW listings Conservation Why uncommon in the GBMWHA Why Restricted distribution – sandstone outcrops between the Central Coast and Putty. Restricted distribution Widespread on shallow soils on Hawkesbury sandstone plateaus near the coast. soils on Hawkesbury on shallow Widespread Uncommon in the GBMWHA. Widespread on moderately fertile soils drier parts of the coastal plain and escarpment Widespread Its preferred combination of soil fertility and climate is Victoria. from Queensland to limited in the GBMWHA. Widespread along the western slopes north from Denman. Suitable climatic conditions Widespread north of the GBMWHA. only occur in the far Restricted distribution – sandstone ridges around northern Wollemi NP, Goulburn River River Goulburn NP, Wollemi – sandstone ridges around northern Restricted distribution NP and Manobolai NR Occurs on cold alluvial flats from Wallerawang to Victoria. Suitable climatic and to Wallerawang Occurs on cold alluvial flats from drainage conditions are limited in the GBMWHA. Restricted distribution – scattered populations on skeletal soils between Linden and and Linden between soils skeletal on populations scattered – distribution Restricted Berrima Restricted distribution – scattered populations at Wentworth Falls, Budawang Range, Budawang Falls, Wentworth – scattered populations at Restricted distribution NP. Wadbilliga Alluvial soils along streams from Putty south to Victoria. Suitable fertile alluvial soils are Victoria. Alluvial soils along streams from Putty south to limited within the GBMWHA. A species endemic to the GBMWHA, only found on the Wollemi and Hunter Ranges. Wollemi A species endemic to the GBMWHA, only found on Restricted distribution – alluvial soils in the lower Hawkesbury-Nepean catchment Hawkesbury-Nepean – alluvial soils in the lower Restricted distribution Locally frequent in drier areas between Narrabri and Nowra. Suitable soil and climatic Locally frequent in drier areas between Narrabri and Nowra. conditions are limited in the GBMWHA. A species endemic to the GBMWHA, with scattered populations on skeletal soils at A species endemic to the GBMWHA, with scattered populations on skeletal elevations. lower Widespread along the western slopes north from about the Goulburn River into River along the western slopes north from about Goulburn Widespread north of the GBMWHA. Queensland. Suitable climatic conditions only occur in the far On open swampy flats from Nullo Mountain to the Megalong Valley. Suitable swampy Suitable swampy Valley. flats from Nullo Mountain to the Megalong On open swampy alluvial soils are limited within the GBMWHA. Restricted distribution – Bylong to the upper Wolgan Valley on moderately fertile soils. Valley Wolgan – Bylong to the upper Restricted distribution are uncommon in the GBMWHA. Suitably fertile soils within its limited distribution Locally frequent on sandy soils on coastal foothills between Karuah and Nerriga. Locally frequent on sandy soils coastal foothills between Karuah and Nerriga. Uncommon in the GBMWHA. Widespread along the western slopes from Wagga Wagga to Queensland. Its preferred Wagga Wagga along the western slopes from Widespread limited parts of combination of a dry climate and fertile soils only occurs in very Wollemi. northern Restricted distribution – upper Cudgegong River on sandstone ridges. River – upper Cudgegong Restricted distribution A species endemic to the GBMWHA, with localised populations on skeletal soils in the A species endemic to the GBMWHA, with localised populations on skeletal Tableland. Wanganderry upper Blue Mountains and GBMWHA reserves Yengo, Wollemi Yengo, Yengo, Wollemi Yengo, Wollemi Wollemi Wollemi Blue Mtns Blue Mtns, Nattai Blue Mtns Wollemi Wollemi Blue Mtns, Nattai Wollemi Wollemi, Blue Mtns, Nattai Wollemi, Wollemi Wollemi, Blue Mtns Wollemi, Wollemi, Gardens of Stone Wollemi, Yengo Wollemi Wollemi Blue Mtns, Kanangra Nattai Boyd, Eucalypt species euryphylla Angophora Angophora hispida Angophora Corymbia maculata Corymbia trachyphloia subsp . Corymbia trachyphloia amphistomatica Eucalyptus aenea Eucalyptus aggregata Eucalyptus apiculata Eucalyptus baeuerlenii Eucalyptus baueriana Eucalyptus bensonii Eucalyptus benthamii Eucalyptus beyeriana Eucalyptus burgessiana Eucalyptus caleyi subsp . caleyi Eucalyptus caleyi Eucalyptus camphora subsp . Eucalyptus camphora camphora Eucalyptus cannonii Eucalyptus capitellata Eucalyptus conica Eucalyptus corticosa Eucalyptus cunninghamii on overall distribution are from PlantNet and Fairley and Moore (2000). are from PlantNet and Fairley distribution on overall Table 1b. Eucalypt species with relatively restricted distributions in the Greater Blue Mountains World Heritage Area showing occurrence in conservation reserves, comments on in reserves, showing occurrence conservation Area Heritage in World distributions Eucalypt the Blue restricted species Greater Mountains with 1b. relatively Table listings. conservation informal and formal and distribution, overall the Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 429

3RCa LC * 2V 3RC- LC *

3RCa

2RC- LC *

2E 3KC DD * 3RCa 2ECi CR D1* Vulnerable Ecological Vulnerable Community (TSC) Vulnerable (TSC) Vulnerable Vulnerable (TSC) Vulnerable Endangered (TSC) Endangered (TSC) Endangered (EPBC) Sufficiently cold conditions are Victoria. Locally frequent on the western slopes and Hunter valley between Wellington and Wellington between Locally frequent on the western slopes and Hunter valley The suitable combination of climate and soil fertility only occurs on Singleton. Wollemi. escarpment footslopes in northern Restricted distribution – scattered populations from Mt Tomah to the Budawang Range. to the Budawang Tomah – scattered populations from Mt Restricted distribution Widespread on shallow soils along the western slopes from Victoria to Queensland. Victoria soils along the western slopes from on shallow Widespread north of the GBMWHA. Suitable climatic conditions only occur in the far Restricted distribution – Lake Macquarie and northern Yengo NP to Mountain Lagoon Yengo Macquarie and northern – Lake Restricted distribution Restricted distribution – sandstone ranges between the Hunter valley and northern Yengo NP Yengo northern and valley Hunter the between ranges sandstone – distribution Restricted Localised distribution – the lower Hunter Valley to Kangaroo Valley to Kangaroo Valley Hunter – the lower Localised distribution Widespread on the tablelands from Mt Coricudgy to Queensland. A species typical of the on the tablelands from Mt Coricudgy to Queensland. Widespread with a disjunct population south of the Hunter valley Tablelands Northern Restricted distribution – skeletal soils from Newnes Plateau to Nullo Mountain soils from Newnes – skeletal Restricted distribution Disjunct populations on sandy soils between the Gibraltar Range and Deua NP Restricted distribution – cold flats on the Southern Highlands, with an outlying Restricted distribution Plateau occurrence on the Boyd Highly disjunct distribution – Broke to St Albans, Enmore to Wollomombi and in in and Wollomombi to Enmore Albans, St to Broke – distribution disjunct Highly Queensland. Disjunct occurrences on sandy soils in the Gibraltar Range, Blue Mountains & Range Budawang Widespread along the coast and escarpment from Bindook Highlands to Victoria. It Victoria. along the coast and escarpment from Bindook Highlands to Widespread reaches its northern limit in the GBMWHA. On sandy soils along the western slopes north from about Capertee to Queensland. Suitable climatic and soil conditions are limited within the GBMWHA. Restricted distribution – skeletal soils from the Watagans to Colo Heights Watagans soils from the – skeletal Restricted distribution Disjunct occurrences along the escarpment – Bundanoon to Bindook Highlands, to Dorrigo Tops Barrington Restricted distribution – sandstone soils from Lake Burragorang to Yalwal Plateau Yalwal Burragorang to – sandstone soils from Lake Restricted distribution to on cold flats south from Sofala Widespread limited in the GBMWHA. Central Coast to northern Sydney, Colo Heights area; mainly occurs on hilly parts of the Central Coast to northern Sydney, coastal plain, uncommon in the GBMWHA. On sandstone from Cessnock to near Picton. Uncommon in the GBMWHA because it mainly occurs on plateaus nearer the coast. Widespread on cold flats at higher altitudes from the McPherson Range to Victoria. on cold flats at higher altitudes from the McPherson Range to Widespread Sufficiently cold conditions are rare in the GBMWHA. Highly localised at Wentworth Falls (but only recorded just outside the GBMWHA (but Falls Wentworth Highly localised at boundary) reserve Restricted distribution – sheltered gullies in northern Wollemi and Yengo. Awaiting Awaiting Yengo. and Wollemi – sheltered gullies in northern Restricted distribution formal description before inclusion in the GBMWHA list. Sporadic and scattered, in mallee heath on sandy soils of limited drainage in elevated Sporadic and scattered, in mallee heath on sandy soils of limited drainage elevated of GBMWHA). Plateau (on margin to Newnes Wadbilliga areas; Awaiting formal description before inclusion in the GBMWHA list. Awaiting Reported near Mount Coricudgy (Stephen Bell pers. com. 2010). Still to be confirmed. Wollemi Blue Mtns Wollemi Yengo, Wollemi Yengo, Yengo Yengo, Wollemi, Blue Wollemi, Yengo, Mtns, Nattai Wollemi Wollemi, Gardens of Stone Wollemi, Gardens of Stone, Blue Mtns Kanangra Boyd Yengo Blue Mtns, Kanangra Boyd Blue Mtns Wollemi, Gardens of Stone Wollemi, Yengo, Wollemi Yengo, Blue Mtns, Kanangra Boyd Blue Mtns Kanangra Boyd Wollemi Yengo, Wollemi Yengo, Blue Mtns, Kanangra Boyd Not recorded in GBMWHA in recorded Not reserves Yengo, Wollemi Yengo, Not recorded in GBMWHA in recorded Not reserves Wollemi Wollemi Eucalyptus dawsonii Eucalyptus dendromorpha Eucalyptus dwyeri Eucalyptus fergusonii subsp . Eucalyptus fergusonii dorsiventralis Eucalyptus fracta Eucalyptus hypostomatica Eucalyptus laevopinea Eucalyptus laophila Eucalyptus ligustrina Eucalyptus macarthurii Eucalyptus michaeliana Eucalyptus moorei Eucalyptus muelleriana Eucalyptus nubila Eucalyptus prominula Eucalyptus quadrangulata Eucalyptus ralla Eucalyptus rubida subsp . Eucalyptus scias subsp. Eucalyptus squamosa Eucalyptus stellulata Potential additional species Potential Eucalyptus copulans Eucalyptus expressa ms. (also Eucalyptus expressa as Eucalyptus sp. aff. known (Bees Nest Ridge) eugenioides Eucalyptus gregsoniana sp. Howes Swamp Swamp Eucalyptus sp. Howes Creek (Doherty 26) Eucalyptus nobilis 430 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area with records of recent surveys by the New South Wales district on the mid north coast of NSW. Hence, Eucalyptus Department of Environment, Climate Change and Water rudderi has been replaced by Eucalyptus hypostomatica in (DECCW) and its predecessors over the past 20 years. Since the current list. the inscription of the GBMWHA in 2000, vegetation surveys The original nomination included an undescribed taxon, have been undertaken in the Jenolan Karst Conservation Eucalyptus sp. aff. dalrympleana, which has not been included Reserve, Nattai, Thirlmere Lakes, Yengo, Gardens of Stone in the current list because there is insufficient information to and much of Blue Mountains National Park. The survey of attribute it to any newly described or undescribed species. Wollemi NP is currently in progress, and surveys are in the planning stage for Kanangra Boyd NP and the Grose Valley. Subspecies names were not attributed to Eucalyptus radiata A number of eucalypts not known from the GBMWHA and Eucalyptus mannifera in the nomination. We have listed at the time of its inscription have now been confirmed. Eucalyptus radiata subsp. radiata as it is the only subspecies Nomenclature and authorities follow PlantNet, the website occurring in the GBMWHA. The reference to Eucalyptus for the National Herbarium of NSW. (Note: the term species mannifera could have been to either of Eucalyptus mannifera (rather than taxa) is used in this paper in the broadest sense subsp. gullickii or Eucalyptus mannifera subsp. mannifera. and includes subspecific taxa). However, it was probably the former because this taxon is more widespread in the Blue Mountains. 98 recognised eucalypt taxa (mostly species of Eucalyptus but including five Angophora and four Corymbia species) The current list remains subject to change as further survey have been recorded in the eight reserves of the GBMWHA and exploration bring to light new occurrences of species, (Tables 1a and 1b). The species range in size from large forest and perhaps even new species. Bell (2008) in his list of rare trees over 30 m in height (e.g. Eucalyptus deanei, Eucalyptus or threatened plants of Wollemi NP, includes two species saligna), through spreading woodland trees (e.g. Eucalyptus that were not formally described at that time. Eucalyptus sp sclerophylla) to small multistemmed mallees, some less aff. eugenioides (Bees Nest Ridge) is a stringybark found in than 1 m in height in heathland (Eucalyptus cunninghamii). sheltered gullies in northern Wollemi and Yengo NPs (to be More than half of the species (58) (Table 1a) are regarded as named Eucalyptus expressa – Bell, Klaphake & Nicolle, in widespread in the GBMWHA i.e. recorded from two or more prep). Eucalyptus sp. Howes Swamp Creek (Doherty 26) is reserves (e.g. Eucalyptus punctata), but others recorded from only known from one small stand growing on alluvium in only one reserve (42 species) (Table 1b) (e.g. Eucalyptus the Mellong Swamps (and is listed as Endangered under the baeuerlenii), may be relatively to very restricted, although EPBC and TSC Acts). As neither taxon has been formally the great disparity in size of the reserves (Figure 1) makes described (2010) they are included in Table 1b as potential this a very rough measure. The restricted eucalypts are either additional species. widespread species with habitat requirements that are of A small population of mallee-like red gums currently limited extent within the GBMWHA or species that have a attributed to Eucalyptus dealbata has been recorded near naturally restricted distribution. The former are usually more the Martindale Trail in northern Wollemi NP (Bell 2001). typical of drier, colder or more coastal environments. Confirmation of the identity of these specimens is required because this population would represent a considerable Clarification of the list of eucalypts in the original range extension and it occurs on infertile soils on Narrabeen GBMWHA nomination Sandstone, which is an unusual substrate for this species. This taxon has not been included in Table 1b. 90 eucalypts were included in the List of Eucalypts in the GBMWHA nomination (Appendix 6.2 in NPWS 1998). Further change is also likely to result from ongoing However, two of those (Eucalyptus siderophloia & Eucalyptus systematic and ecological research into the relationships umbra) have not been subsequently confirmed as occurring between species. For example recent work on the scribbly within the GBMWHA. Eucalyptus siderophloia is found on gum group (Pfeil & Henwood 2004) indicates that this group moderately fertile soils on the coastal plain but the nearest is best treated as 3 taxa rather than the current 5. If accepted records to the GBMWHA are east of Yengo National Park. and applied to the GBMWHA list this would probably Eucalyptus umbra has also been recorded near Yengo National mean replacing Eucalyptus racemosa, Eucalyptus rossii and Park. While records for both species are close to Yengo NP, no Eucalyptus sclerophylla with Eucalyptus racemosa subsp. confirmed GBMWHA records have been found. racemosa and Eucalyptus racemosa subsp. rossii. Eucalyptus rudderi was included in the nomination because it was formerly considered to occur from Taree to near the Rare or threatened species Shoalhaven River (eg Brooker & Kleinig 1983). Johnson Two thirds of the species have a number of populations in at and Hill (1990) separated Eucalyptus hypostomatica from least two of the eight GBMWHA reserves indicating many Eucalyptus rudderi on the basis of the former’s broad- species are relatively widespread and well-conserved in the lanceolate to ovate, hypostomatic adult leaves. This meant region (Table 1a). that Eucalyptus rudderi is now confined to the Taree-Karuah Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 431

Of those limited to one national park (Table 1b), some are at Eucalyptus macarthurii primarily occur on agricultural the edge of a much wider distribution beyond the GBMWHA. lands with modified pastures. They are threatened by lack (e.g. Eucalyptus dwyeri, Eucalyptus muelleriana). Others of regeneration and are poorly represented in conservation are very restricted. Some such as Eucalyptus bensonii, reserves. Eucalyptus benthamii is threatened by increased Eucalyptus baeuerlenii, and Eucalyptus laophila are nutrients, frequent burning and weed invasion in its habitat very restricted but have most or all of their distribution of alluvial flats. Much of its habitat has been cleared for protected within the GBMWHA. Others (e.g. Eucalyptus agriculture or submerged beneath the waters of Lake hypostomatica, Eucalyptus michaeliana) have reasonably Burragorang (Benson 1985). large populations within the GBMWHA. Six of the rare Eucalyptus dawsonii is a community dominant in the eucalypts are mallees with three being endemic to the Vulnerable Ecological Community Hunter Valley Footslopes GBMWHA (Eucalyptus bensonii, Eucalyptus burgessiana, Slaty Gum Woodland in the Sydney Basin Bioregion that Eucalyptus cunninghamii). occurs below the escarpment in northern Wollemi NP near Other restricted species are less well protected. Six are Jerrys Plains. This community is threatened by clearing, listed as vulnerable under the NSW Threatened Species emerging weeds and frequent burning (Peake 2006). Conservation Act (TSC Act) a low proportion (6%) compared A further ten eucalypts have been recognised as rare on the with listings for other parts of NSW as the extensive area of list of Rare or Threatened Australian Plants (ROTAP, Briggs the GBMWHA has provided conservation security for most & Leigh 1996).). These eucalypts typically have a highly species. Two of these species are also listed as Vulnerable restricted or disjunct distribution, but are subject to a lower under the Commonwealth Environment Protection and level of threat. A good example is Eucalyptus baeuerlenii, Biodiversity Conservation (EPBC) Act. Due to their small which has a very restricted distribution on cliff ledges at population sizes, the primary threat to Eucalyptus fracta Wentworth Falls–Leura, but is within the GBMWHA and is and Eucalyptus corticosa are risks from chance stochastic being managed for conservation. events. Eucalyptus aggregata, Eucalyptus cannonii and

Fig. 2. Many eucalypt species grow in the dissected sandstone landscapes of the Blue Mountains and Wollemi areas where habitats range from moist sheltered gullies to exposed ridges and clifflines. 432 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

Table 2. Classification of eucalypts of the Greater Blue Mountains World Heritage Area showing subgeneric groups and lower groups in Eucalyptus (after Hill 2002) with provisional notes on generalised group ecology.

Genus Generalised group ecology

Angophora Angophora costata Large or small trees on range of soils and rainfall, flower Summer, seeds Angophora floribunda shed at maturity, prolific seedling recruitment Angophora euryphylla Angophora bakeri Angophora hispida Corymbia Corymbia maculata Trees on range of soils and rainfall, mature seeds retained on tree Corymbia gummifera Corymbia eximia Corymbia trachyphloia subsp. amphistomatica Eucalyptus Symphyomyrtus – Adnataria Eucalyptus aenea Mostly woodland or forest trees, mostly Ironbarks or boxes; generally Eucalyptus albens lower rainfall areas, medium nutrient soils clays or clay-influenced Eucalyptus baueriana sandstones; Flower Winter–Spring, seed shed at maturity; recruitment Eucalyptus beyeriana respond to seasonal conditions; long-lived, slow-growing Eucalyptus bosistoana Eucalyptus caleyi subsp. caleyi Eucalyptus crebra Eucalyptus dawsonii Eucalyptus fergusonii subsp. dorsiventralis Eucalyptus fibrosa Eucalyptus hypostomatica Eucalyptus melliodora Eucalyptus moluccana Eucalyptus nubila Eucalyptus paniculata subsp. paniculata Eucalyptus sideroxylon Eucalyptus conica Eucalyptus fracta Eucalyptus polyanthemos subsp. polyanthemos

Eucalyptus Symphyomyrtus – Bisectaria Eucalyptus squamosa Tree or mallee on low nutrient soil, flowers Winter, seed retained for up to a year. Only GBMWHA representative of a group of mainly Western Australian mallees. Eucalyptus Symphyomyrtus – Exsertaria Eucalyptus amplifolia subsp. amplifolia ‘Redgum’ group; Trees on a range of soil and rainfall conditions; flower Eucalyptus blakelyi Spring or Summer, seed retained for up to a year, recruitment episodic, Eucalyptus dwyeri quick growing Eucalyptus michaeliana Eucalyptus parramattensis subsp. parramattensis Eucalyptus tereticornis Eucalyptus Symphyomyrtus – Maidenaria Eucalyptus aggregata ‘Mountain Gums’ on medium or low–medium nutrient soils, often Eucalyptus baeuerlenii montane or cooler Tablelands sites, often with higher rainfall, riparian Eucalyptus benthamii or periodic swamp requirements; Flowering in all seasons depending Eucalyptus bicostata on individual species, seeds retained for a year or shed at maturity Eucalyptus bridgesiana depending on season. Fire response species-specific, some strongly Eucalyptus camphora subsp. camphora epicormic, others weakly epicormic and relatively fire-sensitive. Eucalyptus cypellocarpa Eucalyptus dalrympleana subsp. dalrympleana Eucalyptus mannifera subsp. gullickii Eucalyptus mannifera subsp. mannifera Eucalyptus ovata Eucalyptus quadrangulata Eucalyptus rubida subsp. rubida Eucalyptus smithii Eucalyptus viminalis Eucalyptus macarthurii Eucalyptus praecox Eucalyptus corticosa Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 433

Genus Generalised group ecology

Eucalyptus Symphyomyrtus – Transversaria Eucalyptus deanei ‘Coastal Blue Gums’ , quick-growing forest trees,on medium – high Eucalyptus notabilis nutrient soils, coastal, low elevation sites; mostly Summer flowering Eucalyptus punctata with mature seeds shed sporadically after the next Summer. Generally Eucalyptus resinifera subsp. resinifera vigorous seedling recruitment. Eucalyptus scias subsp. scias Eucalyptus saligna

Eucalyptus Eucalyptus – Blue-leaved ashes A Eucalyptus oreades Woodland or forest trees, often community dominants, on low – high Eucalyptus pauciflora nutrient soils, flowering Summer, seeds retained in capsules for up to a Eucalyptus piperita year, fire response variable, strong epicormic or resprouting to killed (E. oreades). Seedling recruitment vigorous, not fire dependent, often colonisers of disturbed sites. Eucalyptus Eucalyptus – Blue-leaved ashes B Eucalyptus consideniana Mainly woodland trees, often community dominants, on low nutrient Eucalyptus multicaulis soils; flowering Spring/Summer, seeds retained in capsules for up to Eucalyptus rossii a year; fire response strong epicormic or lignotuberous resprouting. Eucalyptus sclerophylla Seedling recruitment sporadic. Eucalyptus sieberi Eucalyptus Eucalyptus – Black sallies Eucalyptus moorei Mallees or small trees, low to medium nutrient soils at high elevation, Eucalyptus stellulata flowering Autumn Eucalyptus Eucalyptus – Green-leaved ashes Eucalyptus apiculata Mainly long-lived mallees, on low-nutrient sandy soils, flowering Eucalyptus burgessiana Summer, with seed retained in capsules for about a year, vigorous Eucalyptus cunninghamii lignotuberous regrowth after fire, seedlings rarely reported. Eucalyptus Eucalyptus dendromorpha fastigata, large tree on high nutrient soils is exception. Eucalyptus fastigata Eucalyptus laophila Eucalyptus obliqua Eucalyptus stricta

Eucalyptus Eucalyptus – Peppermints Eucalyptus dives Trees, on range of soils, flower Spring, seeds shed or retained, juvenile Eucalyptus elata recruitment generally conspicuous, fire response strong to weak Eucalyptus pilularis epicormic growth Eucalyptus radiata subsp. radiata Eucalyptus Eucalyptus – Stringybarks Eucalyptus agglomerata A large group of trees, small trees and mallees often with populations Eucalyptus bensonii intergrading with nearby stringybark species; mostly on low nutrient Eucalyptus blaxlandii soils, but some on medium and high nutrient soils, over a wide range Eucalyptus cannonii of rainfall, though all in well-drained sites. Flower mainly Autumn– Eucalyptus capitellata Winter, sporadically. Seed retained for up to a year, seedling recruitment Eucalyptus eugenioides sporadic. Eucalyptus globoidea Eucalyptus laevopinea Eucalyptus ligustrina Eucalyptus macrorhyncha Eucalyptus muelleriana Eucalyptus prominula Eucalyptus ralla Eucalyptus sparsifolia Eucalyptus tenella 434 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

Significant species just outside the GBMWHA a very wide range of habitats and some groups including Angophora, Corymbia and Symphyomyrtus–Adnataria An endangered species Eucalyptus copulans now persists include species with a very wide range of ecologies. In the as only two individuals in suburban Wentworth Falls within GBMWHA eucalypts are only absent from rainforest and a kilometre of the boundary of the GBMWHA. Because swamp sites. Competition for light at the seedling recruitment of the clearing for urban development it is now too late to stage probably keeps them out of rainforest, as there are many determine its original distribution, but it was evidently very eucalypts that grow on the high nutrient soils that are generally restricted and indicates the vulnerability to human impacts associated with rainforest. Most eucalypts need well-drained of species with very restricted occurrences. soils and are killed by extended waterlogging. However some Eucalyptus gregsoniana which has a ROTAP code of species in Symphyomyrtus–Maidenaria do cope with some 3RCa occurs on Newnes Plateau in close proximity to the waterlogging (Eucalyptus camphora, Eucalyptus aggregata, GBMWHA. It typically occurs above 1100m elevation and Eucalyptus ovata) and could allow evolution of future swamp may be present in the higher parts of the catchments of the species. Wollangambe River, the Wolgan River or Carne Creek. In terms of Grime’s (2001) CSR triangle of ecological strategies COMPETITORS — STRESS — TOLERATORS Classification of GBMWHA eucalypt species — RUDERALS, groups such as the Eucalyptus–Green-leaved ashes and Symphyomyrtus–Adnataria are very strong stress- The genus Eucalyptus was described in 1792 from specimens tolerators, while Symphyomyrtus–Transversaria are strong of Eucalyptus obliqua from Tasmania. However specimens of competitors. The majority of species have components of both Angophora costata and Corymbia gummifera were collected strategies however and this has probably contributed to their by Joseph Banks and Daniel Solander at Botany Bay in 1770 success. Ruderal strategy characteristics (short-life, quick and named by Joseph Gaertner in 1788 as Metrosideros growth, colonisers) show in a couple of species from different costata and Metrosideros gummifera respectively. The first subgroups (Eucalyptus oreades, Eucalyptus elata) but are eucalypt collections from the GBMWHA were probably unusual. These strategies may provide some pictures of past made by the botanist George Caley who explored Thirlmere dynamic changes. For example the stress-tolerator capacity Lakes in 1802 and the lower Blue Mountains reaching as far of the Eucalyptus–Green-leaved ashes and their geographical as Mount Banks in 1804. distribution on ridges particularly in the upper Blue Mountains Description of the 700 or so eucalypts has involved many suggest they may have been able to survive in that area during prominent Australian botanists including Ferdinand Mueller, the cold conditions of the Last Glacial Maximum 20 000 years Joseph Maiden and William Blakely. In 1971 Pryor and ago, whereas competitor groups such as Symphyomyrtus– Johnson (1971) proposed a classification dividing the Transversaria may have been able to invade new areas as group into a number of subgenera; subsequent research climate ameliorated 5–10 000 years ago. has supported the various subgeneric groupings, but raised questions over their relative ranks. For the eucalypts of the Habitat and ecological ranges of GBMWHA eucalypts GBMWHA three genera are currently recognised, Angophora, Corymbia (raised to generic rank by Hill & Johnson 1995) Brief notes on the habitats of the individual species (Table and Eucalyptus, which itself includes two major subgenera 3) have been compiled predominantly from DECCW Eucalyptus (Monocalyptus) and Symphyomyrtus. Further surveys (NPWS 2003, DEC 2004, DEC 2006, DEC 2006a & subdivision of these subgenera into the formal and informal DECC 2008). Sources of information for those parts of the groups (following Hill 2002) is given in Table 2. GBMWHA where survey is not complete were Bell (1998) for Wollemi National Park, Keith and Benson (1988) and Ecological strategies Black (1982) for Kanangra Boyd National Park and Keith and Benson (1988) and Benson (1992) for the Grose valley. While many of the subgeneric groups align with groups based The main landscape characteristic of the GBMWHA is the on bark characters that have been traditionally used in field predominance of sandstone but within this is a considerable identification and forestry work particularly, e.g. stringybarks, variation in local habitat conditions resulting from boxes, ironbarks, others cut across several groups e.g smooth- topographic variation, climate, slope and aspect, and soil barked gums. However it is considered that further insights into fertility. The following factors contribute to the diverse range the identification and ecology of the individual species might of habitats present in the GBMWHA. be gained from consideration of common attributes relevant to the groups. With this in mind, general ecological features Altitude and climate that appear to be characteristic of particular groups, including soil nutrient conditions, habitat, flowering and fruiting timing Elevation in the GBMWHA ranges from 20 m elevation and fire responses have been listed using data sourced from near the Nepean River at Glenbrook to 1 330 m elevation at Benson & McDougall (1998) together with our general field Mt Emperor near Jenolan Caves. The lowest plateau in the observations on the species. Eucalypts have evolved to occupy GBMWHA is Yengo NP which has extensive areas below Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 435 Wollemi area Valley Burragorang and around Lake Valleys Burragorang Range and around Lake Tonalli and Howes Wallabadah Valleys Valleys Valleys, the Blue Labyrinth & Nattai Wollemi, Valley & Yango and Yengo Nattai, Little River Wollondilly, Blue Labyrinth, Bonnum Pic Tableland, Wollemi Wollemi and Scotts Main Range Valley catchment River areas of the Goulburn rainfall 400 m and the low below Wollemi Wollondilly and tributaries River Wollondilly area to Martindale Range (localised) Yengo (localised) Wollemi limited) Range between Cyrils Rocks and Gospers Mtn (very Wollemi on sandstone and with thin layers of shale, the Blue Labyrinth, Breaks in limited parts south east Yengo on Jamison Ck Walk Habitat/distribution the escarpments of colluvial slopes on Permian sediments below • dry, 400 m in southern slopes on sandstone below • ridges, upper slopes and lower ridges and plains with deep, freely draining sands on the Mellong Plateau • low 700 m situations on dissected sandstone plateaus across the GBMWHA below • sheltered slopes and gullies more exposed and Kedumba • drier slopes on Permian sediments beneath the escarpment in Megalong in the Erskine Ranges diatremes at mid elevations volcanic • larger • sandstone outcrops from Mt of substrates throughout the GBMWHA creeks and gullies on a variety larger • alluvium along major rivers, • colluvial slopes beneath the escarpment across GBMWHA area • well drained sandy alluvium of the Mellong Swamps Plateau and Newnes pagodas between Dunns Swamp • more limited among rocky (localised) ridges north of Upper Colo and near Mt Lockyer • exposed • throughout 700 m and sheltered slopes on dissected sandstone plateaus across the GBMWHA below • more exposed and Kedumba • drier slopes on Permian sediments beneath the escarpment in Megalong • sandstone ridges and sheltered situations enriched by shale in a small area near Bulga areas of northern rainfall aspects on sandstone in low • exposed • dry ridges in north western • the Bindook Highlands, steep slopes of Flat, Murruin Range Limeburners and alluvial bogs at high elevations, • moist alluvial soils associated with cold air drainage hollows areas north of the Hunter Range rainfall areas in low rainfall • basalt caps and diatremes in low slopes and drainage lines in the Capertee siltstone & limestone on lower • Devonian • undulating foothills along the across the GBMWHA and sheltered slopes on dissected sandstone plateaus at moderate elevations • more exposed Nattai, Little and Burragorang the escarpment of Kedumba, • dry colluvial slopes below of streams in Big at the headwaters • alluvial flats along streams and colluvial fans Blue Mountains and eastern diatremes of the Lower • volcanic Linden, Kings 900m, eg sandy soils below • rock plates with skeletal • Darwins • alluvial flats along Putty Creek soils along the • skeletal catchments Creek, Reedy Little and Nattai River • alluvial flats in the Kedumba soils in eastern slopes with shallow • ridges and exposed areas north of Nullo Mountain rainfall • basalt caps in low • limited – Karst at Jenolan Caves Mt Nullo Mountain, Cedar Creek areas, eg • high altitude >900m basalt caps in rainfall on Gangerang & Murruin Range • sheltered gullies and protected slopes at higher elevations Eucalypt species bakeri Angophora Angophora floribunda Angophora Angophora costata Angophora euryphylla Angophora hispida Angophora Corymbia eximia Corymbia gummifera Corymbia trachyphloia subsp. Corymbia trachyphloia amphistomatica Corymbia maculata Eucalyptus albens Eucalyptus aenea Eucalyptus agglomerata Eucalyptus aggregata Eucalyptus amplifolia subsp. amplifolia Eucalyptus apiculata Eucalyptus baueriana Eucalyptus bensonii Eucalyptus baeuerlenii Eucalyptus benthamii Eucalyptus beyeriana Eucalyptus bicostata Table 3. Notes on the habitats of individual Greater Blue Mountains World Heritage Area eucalypt species compiled predominantly from DECCW survey work. work. DECCW survey from eucalypt species compiled predominantly Area Heritage World Blue Mountains Greater 3. Notes on the habitats of individual Table 436 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area Valley

Valleys

Wollondilly

Burragorang

the

&

of

Nattai

slopes

the

to

steep

and

Valleys

Highlands Jamison

&

Valleys Bindook

Kedumba

the Blue Mountains ridge, the Lower Golden Moon Bluff Tableland, Range,

Wollondilly

St Helena Crater The Ironbarks, Euroka, Machins Crater, Wollemi from

Murruin

Wollemi to

benches

eg Wollemi, and Yengo and

Caves

Plateau Burragorang and Newnes arm of Lake High Gangerangs, Coxs River Valleys, Leap and Mt Banks Govetts Narrowneck, Falls, Wentworth Jenolan

midslopes Mountain, Lacys Axehead

& Barton Creeks Crown eg Valleys, & Barton Creeks Crown eg Valleys, and

Valley Valley Walls, Range

Wolgan Wolgan Neck Narrow below Valley escarpment

River Wollondilly Black

Wollondilly Wollondilly protected from

on

& upper Grose plateaus near Jamison valley Tableland, elevations

sediments

(localised) Wollemi 800 m, and Blue Labyrinth south of the Hunter Range below Wollemi high

at

Permian

(currently outside GBMWHA), Caves Wombeyan on gullies

and soils

on Permian sediments Valley slopes

colluvial

sheltered deeper

Habitat/distribution areas of north west rainfall sandy soils in sheltered situations the low • shallow areas north of Nullo Mountain rainfall • diatremes in low on the floor of Capertee & • undulating topography Gangerang Range to metasediments between 800 and 1000m, Black Range, Jenolan Caves, Silurian and Devonian • sheltered slopes & ridges on Ordovician, trail; Limeburners • Bindook Highlands and the steep slopes of Budthingeroo, Mt Cameron areas, eg • high altitude >900 m basalt caps in rainfall sheltered slopes and ridges with deeper soils on sandstone pagoda complexes; • gullies among rocky at Mt Jellore (limited) • on trachyte • Bindook Highlands and the steep slopes of tributaries and larger Rivers • occasional on sheltered aspects along the Jenolan, Coxs and Kowmung slopes around • rocky Box Hole Clearing 500 m, eg areas above • higher altitude diatremes in high rainfall (limited) River • sandy soils along creeks around the Cudgegong • slopes along Murruin Creek, Jocks Creek and the Flat, Murruin Range Limeburners and alluvial bogs at high elevations, • cold air drainage hollows Kedumba 900 m, eg soils below • rock plates with skeletal catchment River areas in the Goulburn rainfall • ridges and slopes in low soils on the footslopes of Megalong • waterlogged & Ganguddy Creeks Never • poorly drained alluvial sands possibly associated with impermeable clays in Never Gulph and the Capertee valley the escarpment and protected gullies between Growee • talus slopes below on the floor of Capertee & • undulating topography (localised) • Near the Macdonald River • Fertile soils in north western slopes in • ridges and exposed (localised) River sandstone soils on ridges around the Cudgegong • shallow Nattai, Little and Burragorang valleys • dry & sheltered slopes through the Coxs, Jenolan, Kowmung, and Burragorang between the Kedumba the escarpment along Lake • colluvial slopes below areas of GBMWHA metasediments in higher rainfall • deep, protected gullies and sheltered slopes on sandstone Silurian Devonian Blue Mountains diatremes of the lower • volcanic sandstone plateaus e.g., Kings tableland, • localised on higher elevation Plateau undulating terrain the Boyd 900m at Jenolan Caves, sites above • exposed Plateau of creeks on the Newnes • poorly drained headwaters pagodas across the GBMWHA • protected sandstone gullies, sheltered slopes and among rocky Jamison and Kedumba • slopes beneath the escarpment in Megalong, Plateau to Murruin Range and the Bindook Highlands and Boyd from Black Range, Jenolan Caves • moist, sheltered situations at high elevations Plateau • ridges and sheltered slopes on around Newnes Gulph and Jerrys Plains areas between Growee rainfall the escarpment in low • talus slopes below • • • Capertee soils on Kings • rock plates with skeletal • upper slopes and undulating ridges along Scotts Main Range the Bindook Highlands • ridges and protected slopes on sandstone with interbedded layers of shale in Blue Mountains and eastern diatremes of the Lower • shale caps and volcanic Eucalyptus blakelyi Eucalypt species Eucalyptus blaxlandii Eucalyptus bosistoana Eucalyptus bridgesiana Eucalyptus burgessiana Eucalyptus camphora subsp. Eucalyptus camphora camphora Eucalyptus caleyi subsp. caleyi Eucalyptus caleyi Eucalyptus cannonii Eucalyptus conica Eucalyptus crebra Eucalyptus capitellata Eucalyptus consideniana Eucalyptus corticosa Eucalyptus dendromorpha Eucalyptus dives Eucalyptus dawsonii Eucalyptus deanei Eucalyptus dalrympleana subsp. dalrympleana Eucalyptus cypellocarpa Eucalyptus cunninghamii Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 437 Valley Wanganderry Wollondilly Valleys Plateau Newnes and below Valleys, Wollondilly Valley Valleys Yengo Yengo Wollemi Creek Deanes & Rocky Plateaus, eg and Newnes Ti-Willa Wollondilly Capertee valley Tableland, Ridge Wattle Wollondilly Kings Walls, Hole area Creek & Devils Werong and eastern Yengo Valleys Tablelands; the Blue Breaks and Nattai Wollemi, Wallabadah, between Joorilands and Murruin Creek Valley Yengo, Yango, Wanganderry The Ironbarks River Wollondilly Wollemi < 800 m at elevations Wollemi & Barton Creeks (limited) Crown eg Valleys, Culoul Range, Blue Labyrinth & Nattai, eg Wollemi, Hole Creek, Devils Werong Wolgan areas of northern rainfall aspects in low • exposed • sheltered situations on Murruin Range, Bindook Highlands and steep slopes of the and Kedumba • escarpment footslopes on Permian sediments in Burragorang, Nattai, Megalong • sheltered positions on sandstone across the Nattai and catchments • alluvial flats along major streams in the Little and Nattai River Nattai, Little and Burragorang catchments • dry & sheltered sites through the Coxs, Jenolan, Kowmung, and Burragorang betweeen the Megalong the escarpment on both sides of Lake • colluvial slopes below Blue Mountains • residual shale caps and interbanded soils on sandstone ridges of the Lower in gullies and diatremes Big • localised colluvial fans • upper slopes and undulating ridges along Scotts Main Range the Bindook Highlands steep protected of Plateau to Murruin Range and the Bindook Highlands & Boyd from Black Range, Jenolan Caves • sheltered locations at the high elevations Plateau escarpment on Newnes • gullies & sheltered slopes below Budthingeroo • high altitude basalt caps (limited), eg • enriched soils on gentle slopes, ridges and benches in central northern and Burragorang between the Kedumba the escarpment along Lake • colluvial slopes below • ridges and slopes on sandstone or soils enriched by shale in Tablelands and Burragorang • higher shale caps in south east Rivers of the Coxs & Kowmung • scattered occurrences on sheltered sites in the tributaries Nullo Mtn Mt Coricudgy, areas, eg • high altitude >900m basalt caps in rainfall • between sandstone pagodas in western Kedumba 900m, eg sandy soils below • localised on rock plates with skeletal and Scotts Main Range Rivers • slopes of the Jenolan, Coxs and Kowmung Blue Mountains, eg • shale caps on ridges of the Lower on Permain sediments • ridges and upper slopes within the Capertee valley • Sentry Box Point near Milbrodale (localised) upper slopes of the Blue Breaks, southern Labyrinth, Nattai, • sandstone ridges enriched with shale bands and exposed slopes on Narrabeen sediments across central and northern south and east facing • mid to lower • upper slopes and crests along southern Scotts Main Range the Bindook Highlands escarpment in the Burragorang and Nattai • slopes and benches below Neck of creeks on sandstone the Bindook Highlands, Narrow • poorly drained headwaters catchment relief situations in the upper Cudgegong • low trail positions on metasediments slopes between 800 and 1000m, Gangerang Range to Limeburners • moderately sheltered, rocky in the Burragorang • foothills and slopes on Bindook Porphyry and Scotts Main Range Rivers sediments in the Jenolan, Coxs and Kowmung • slopes on Devonian • localised diatremes, eg • hills of the Capertee & Plateau (localised) • Boyd Gulph and the Capertee valley areas between Growee the escarpment in higher elevation • Permian talus slopes below Crown metasediments west of Pantoneys soils on Ordovician ridges and slopes with skeletal • exposed • slopes along Murruin Creek, Jocks Creek and the slopes on Narrabeen sediments across central and northern south and east facing • mid to lower Eucalyptus dwyeri Eucalyptus elata Eucalyptus eugenioides Eucalyptus fergusonii subsp. Eucalyptus fergusonii dorsiventralis Eucalyptus fastigata Eucalyptus fibrosa Eucalyptus laevopinea Eucalyptus laophila Eucalyptus ligustrina Eucalyptus fracta Eucalyptus globoidea Eucalyptus hypostomatica Eucalyptus mannifera subsp. Eucalyptus mannifera mannifera Eucalyptus melliodora Eucalyptus mannifera subsp. Eucalyptus mannifera gullickii Eucalyptus macrorhyncha Eucalyptus macarthurii Eucalyptus michaeliana 438 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area substrates

of

variety

a

Trig on

Highlands

Valley Bindook

the Wollemi

to Wollondilly

Yengo Nattai and the lower and tributatries River Wollondilly Wollemi Range

Plateau Range, southern Blue Labyrinth and Newnes Tonalli Murruin Plateau and Newnes Falls Wentworth and Yengo

Plateau, Gangerang Plateau Willa and eastern Yengo, Sandy Camp, Kerrabee/Woodlands Yengo, to

Ti The Ironbarks, Culoul Range in Trail in Valley Creek, Pumpkin Hill (limited) Waterfall Plateau

Wollemi and Wareng Boyd

eg Wollemi, Mountain Creek to Nyanga Tomat Valleys area and Mellong Swamps Valley and

Random, Back and Colong Swamp Tomat, Neck, Narrow Walls, Caves

western Tableland, Jenolan Mounts eg Wollemi,

Tablelands (limited) Thirlmere Lakes Range,

Culoul Range (limited) eg Wollemi, Valley and Yengo escarpment cliffs Plateau, often above and Newnes Tableland Black

and Capertee valleys Wolgan Wanganderry from

and Erskine Creek River Warragamba elevations

high

at

House Creek (limited) Watch situations

and the Bindook Highlands (limited) Yerranderie sheltered

areas of rainfall • basalt caps diatremes in low Habitat/distribution along the • undulating hills and escarpment footslopes on Permian sediments Bindook Porphyry Valleys & Kedumba protected slopes on Narrabeen sediments and shale capped ridges in central northern • mid to lower the floor of Hunter • steep, dry escarpment slopes on Permian sediments above Creek, Capertee • on limestone around Crown catchments areas in the Coxs & Kowmung • rainshadow Kanangra greater than 700m, eg sandy soils at elevations • skeletal Heights, Bullnigang • moderately deep soils on the Bindook Highlands, eg Kings 900m, eg sandy soils below • rock plates with skeletal • interbanded and residual shale soils on sandstone ridges of the Blue Labyrinth Breaks catchment (limited) River areas in the Goulburn rainfall • ridges and slopes in low • hills near from Black Range, to Murruin Range the Bindook Highlands • moist, sheltered slopes and gullies of the high elevations • sheltered slopes and gullies between Kings • sheltered positions on the Nattai and the Bindook Highlands, eg • tableland soaks associated with swamps Plateau and at upper Martins Creek • along gently sloping drainage lines on the of Boyd Blue Mountains and south east • residual shale caps on ridges of the Lower • sandy drainage flats adjoining the eastern side of and permanently wet sites on deep sands in the Garland • better-drained (limited) creeks of the western Capertee valley • colluvial soils along the larger Plateau and around Jenolan Caves on granite the Boyd • on undulating terrain or in cold hollows trail on the Loombah Plateau, Bindook Highlands, Murruin Range and Mt Fatigue alluvial bogs and rock plates at high elevations • cold air drainage hollows, • enriched sandstones between the Hole (limited), Dunns Swamp • north of Rowans ridges and upper slopes on sandstone south of the Old Settlers exposed • dry rocky Green Hill, • shale influenced ridges, slopes and gullies of the Oakdale Plateau, eg northern areas rainfall the low • slopes, gullies and ridges with deeper soils on sandstones across much of the GBMWHA, except escarpment of the • residual Permian outcrops on Scotts Main Range; colluvial slopes below soil on residual plateaus or ridges, High Gangerangs to Black Range • rocky the escarpment in • talus slopes below of substrates, in wetter and dry areas all but • possibly the most frequently recorded and widespread Eucalypt of GBMWHA, occurring on a variety the coldest sites breccia, shale, Permian sediments, volcanic metasediments, Bindook Porphyry, • occurs on ridges, slopes, gullies and talus sandstone, Silurian Devonian granite, alluvium and colluvium. • between Rivers of the Coxs & Kowmung • scattered occurrences on sheltered sites in tributaries • areas • high altitude >900m basalt caps in rainfall and Burragorang betweeen the Kedumba the escarpment on both sides of Lake • dry colluvial slopes below southern escarpment from Katoomba to • small sheltered drainage lines on sandstone above and Kedumba • sheltered slopes beneath the escarpments in Megalong • broad flat to gently sloping sites on plateaus north of Oakdale, southern Blue Labyrinth and Breaks • higher ridgelines with shale caps in south east radiata subsp. Eucalyptus moluccana Eucalypt species Eucalyptus moorei Eucalyptus muelleriana Eucalyptus multicaulis Eucalyptus obliqua Eucalyptus nubila Eucalyptus oreades Eucalyptus notabilis Eucalyptus ovata Eucalyptus paniculata subsp. paniculata Eucalyptus pauciflora Eucalyptus pilularis Eucalyptus prominula Eucalyptus parramattensis subsp. Eucalyptus parramattensis parramattensis Eucalyptus praecox Eucalyptus polyanthemos subsp. polyanthemos Eucalyptus piperita Eucalyptus quadrangulata radiata Eucalyptus Eucalyptus punctata Eucalyptus ralla Eucalyptus resinifera subsp. Eucalyptus resinifera resinifera Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 439 Rivers

Axehead Little

&

Walls, Nattai

Tableland, the

Yengo along Wollemi

and

Wanganderry and Wareng Valley

Random, Back, Colong and Bent Tomat, Howes Valleys

Plateau, Creek, Kanangra area, Boyd Wombeyan around

Mounts eg Wollemi, Plateau, Gangerang Kedumba Willa streams

Ti Yengo broader

and the Capertee valley Wollemi Nattai and Mountain, Lacys, Tonalli and north of Upper Colo in south east Yengo along

flats

and north east Yengo Yengo Neck, Narrow Walls, alluvial

and Saint Helena Crater Euroka, Machins Crater, 400 m, eg below Wollemi

area and Mellong Swamps Valley Valley area Wallabadah on Permian sediments Valleys footslopes

and area and Mellong Swamps Valley

& Yango Wollondilly Culoul Range usually on soils enriched by shale or basalt, eg Wollemi, area,

Gorge Wollondilly Tablelands on Bindook Porphyry & tributaries River Wollondilly Swamps

Mellong

Wanganderry the

in

alluvium

sandy

moist metasediments in western soils on Ordovician 800 m on ridges with shallow • below Mountain Plateau and Donkey escarpment of Newnes • talus slopes below pagodas among rocky • sheltered slopes and valleys Plateau on granite the Boyd • on undulating terrain or in cold hollows slopes on Narrabeen sediments mainly in northern • protected gullies and sheltered lower of gullies in Big at the headwaters • colluvial fans slopes in eastern • sheltered gullies and lower • Culoul Range (localised) Neck, Narrow 600m, eg slopes on sandstone plateaus above • ridgetops and exposed and wetter sites on deep sands in the Garland • better-drained catchment River areas in the Goulburn rainfall • ridges and slopes in low 550 m below valleys River Kowmung dry slopes along the Coxs, Jenolan and lower • exposed plateaus, ridges and slopes between 800 950m, Bindook Highlands, north from Little • high elevation High Gangerang, Black & Moorara Range, Jenolan Caves slopes on sandstone, south from the Capertee valley areas, ridgetops and exposed • rocky • residual Permian outcrops on Scotts Main Range • slopes of the Nattai and areas of the south-eastern corner and higher rainfall elevation • lower & Burragorang • Permian outcrops on Scotts Main Range and footslopes around the edge of Megalong sites from Bindook Highlands to Black Range • sheltered and more exposed 800 m • slopes, gullies and ridges on sandstone enriched shale soils across the GBMWHA mostly below Mountain Plateau and Donkey escarpment of Newnes • Permian outcrops on Scotts Main Range and talus slopes below soil on residual plateaus or ridges, High Gangerangs, Black Range and Jenolan Caves often rocky • skeletal, & Jamison • footslopes around the edge of Megalong sites on deep sands in the Garland • better-drained of the • Bindook Highlands and northern fall valleys River Kowmung • slopes along the Coxs, Jenolan and lower areas of the south-eastern and higher rainfall elevation sites in the lower • exposed Flat, Murruin Range, Plateau, Limeburners Boyd and alluvial bogs at high elevations, • cold air drainage hollows Hook Swamps Kanangra eg sandy soils at higher elevations, • rock plates with skeletal • basalt caps, shale capped ridges and enriched mid slopes in north-western Mountain, Loombah Plateau, along the Murruin Range and Mt Fatigue trail Mountain, Loombah Plateau, along the Murruin Range and Mt Fatigue Gap Heffrons catchment, eg & Cudgegong between the Growee • along the divide floors, ridges and upper slopes within the Capertee valley • broad valley and along Scotts Main Range tributaries their larger Rivers, sediments in the Jenolan, Coxs and Kowmung • sheltered & dryer slopes on Devonian Burragorang, Nattai and Little Burragorang, the Kedumba, arm of Lake escarpment on protected aspects in the Coxs River • deeper colluvial soils below Valleys; River • slopes & undulating foothills of the Blue Mountains and south east diatremes of the lower • volcanic • • Bindook Highlands and protected slopes of the Plateau on granite, especially around eastern & northern plateau rim • on gentle slopes across the Boyd Budthingeroo areas, eg • high altitude basalt caps and diatremes in rainfall & the Black Range to Murruin from Jenolan Caves • sheltered sites at high elevations • freely drained alluvial soils along drainage lines on the Mellong sandmass and sheltered streams Plateau and Newnes pagodas between Dunns Swamp streams and among rocky • along larger Eucalyptus rubida subsp. Eucalyptus saligna Eucalyptus rossii Eucalyptus sclerophylla Eucalyptus scias subsp. Eucalyptus sieberi Eucalyptus sideroxylon Eucalyptus smithii Eucalyptus sparsifolia Eucalyptus squamosa Eucalyptus stricta Eucalyptus stellulata Eucalyptus tenella Eucalyptus tereticornis Eucalyptus viminalis 440 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

300 m elevation. The remainder of the GBMWHA is usually mm in the central Blue Mountains (Katoomba to Newnes higher than 400 m elevation, other than the larger incised Plateau) and in Kanangra – Boyd NP. Rainfall progressively valleys such as the Coxs and Grose Rivers. Only a relatively decreases north of Newnes Plateau and is lowest in the small part of the GBMWHA is above 1 000 m elevation, northwest of the GBMWHA, in northern Wollemi. Jerrys mainly around Mount Werong, the Boyd Plateau and Narrow Plains near the northern boundary of Wollemi NP has an Neck Peninsula. A notable feature of the GBMWHA is the annual rainfall 640 mm, which is probably representative high level of vertical relief between the plateaus and incised of this part of the World Heritage area. Taxa that are more valleys, which reaches a peak of about 900 m in the south widespread on the NSW Western Slopes such as Corymbia west of the area. trachyphloia subsp. amphistomatica, Eucalyptus dawsonii and Eucalyptus beyeriana only occur in this part of the As the altitude increases, the climate becomes colder and GBMWHA. wetter. Frost, fog and some snow can occur in the Upper Mountains, where conditions are generally colder and harsher. Species restricted to the coldest areas include Rock types Eucalyptus pauciflora Eucalyptus laevopinea and on the The oldest rocks of the GBMWHA are of Siluro-Devonian Eucalyptus rubida, Eucalyptus stellulata high peaks, and and age and are mainly found in the southern Blue Mountains. Eucalyptus aggregata on flats subject to cold air drainage. Tectonic plate movements and volcanic intrusions have The climate is much warmer in the lower eastern and northern caused considerable deformation to them. Deposition of the areas; those species typically found at lower elevations such Sydney Basin sediments commenced in the early Permian Angophora costata Corymbia eximia as and usually have a with marine sediments and coal deposits which outcrop widespread distribution within the GBMWHA. below the major escarpments. The sandstones that are so extensive across the region were deposited next, as river Rainfall sediments in late Permian to mid Triassic. More fertile shales were occasionally deposited above these sandstones. Annual rainfall over most of the GBMWHA is moderate The most recent phase were the basalt flows of the Miocene, (800–1200 mm pa) but increases with elevation to 1400 which remain as caps on a number of mountains.

Fig. 3. Smooth straight trunks of Eucalyptus deanei in Blue Gum Forest in Blue Mountains National Park Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 441

Through the processes of weathering, rock breaks down into Drainage soil – some rocks create more fertile, higher nutrient soils than others. The sandstones break down into low nutrient soils The often steep topography in the sandstone areas generally which hold little moisture. There are many eucalypts typical allows water to run off rapidly but where there are layers of of low nutrient sandstones including Angophora bakeri, impervious rock, such as claystone or ironstone, waterlogged Angophora hispida, Corymbia gummifera, and many of the conditions may occur allowing swamp communities to subgenus Eucalyptus such as the Eucalyptus–Green-leaved develop. Most eucalypt species are sensitive to waterlogged ashes and Eucalyptus Blue-leaved ashes B. The quartzites conditions and they rarely occur in poorly-drained sites. An typical of the rugged valley of the mid Coxs River produce exception is Eucalyptus camphora found in swamps in the low nutrient soils, where Eucalyptus sieberi and Eucalyptus Rylstone area. Species in alluvial habitats may recruit in punctata commonly occur. The Permian coal measures, temporarily wet sites after floods e.g. Eucalyptus benthamii, shale and volcanic rocks such as basalt and porphyry form a Eucalyptus elata. brown, loamy soil which is high in nutrients and retains more moisture. Distinctively different plant communities occur on Aspect these soils, dominated by tall eucalypts such as Eucalyptus deanei, Eucalyptus fastigata or Eucalyptus muelleriana. The aspect of the slope determines how much sunlight it gets, Localised areas of basalt may have Eucalyptus saligna or and which winds it is more exposed to. In general, north- Eucalyptus viminalis in higher rainfall areas or Eucalyptus facing slopes are drier and hotter, supporting a different albens and Eucalyptus moluccana in drier areas. Rainforests suite of smaller eucalypts than south-facing slopes. Exposed are largely restricted to the higher nutrient soils in the highest sites associated with cliff-lines in the upper mountains may rainfall areas. frequently have the mallees e.g. Eucalyptus cunninghamii, Eucalyptus stricta, though these sites are also influenced by drainage and slope characteristics.

Fig. 4. Mountain Gum forest in Kanangra-Boyd National Park. Large shaggy trunks of Eucalyptus dalrympleana with smaller trees of Eucalyptus pauciflora, Snow Gum, in background. 442 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

Slope dry areas of northern Wollemi NP are eucalypts that occur primarily on the NSW western slopes such as Corymbia The steeper a slope, the more quickly water will run off, trachyphloia subsp. amphistomatica. The southern part of the taking nutrients with it. The combination of slope and aspect GBMWHA has several eucalypts that are widely distributed has a profound effect on the eucalypt species present. South- in southern NSW and Victoria, including Eucalyptus smithii, facing slopes in the Blue Mountains are generally moister Eucalyptus aggregata and Eucalyptus muelleriana. and steeper than north-facing (Holland et al 1992) and have different eucalypt species, particularly on the lower slopes where moisture can accumulate e.g. Eucalyptus saligna, Eucalypts and climate change Eucalyptus cypellocarpa. On the other hand, north-facing slopes more frequently support species tolerant of stronger Changes in rainfall, evaporation, temperature and carbon sunlight and drier soils such as Eucalypus fibrosa or dioxide levels associated with climate change are likely Eucalyptus crebra. to alter the patterns of mortality, growth and phenology of individual eucalypts. It is likely that different species will Fire, disturbances respond differently to these changes, and that species will vary in their level of tolerance to these changes. Those Fire plays an important role in the growth and development species with a more plastic response may be advantaged over of vegetation communities in Australia. After fire most those with limited tolerance to change. As eucalypt seeds do eucalypt trees regenerate from dormant epicormic buds along not move very far in the landscape, large scale movement the trunk or major branches. In some species particularly of individual eucalypt species in response to changing the mallees, the canopy and stems are killed and the plants conditions is not likely, though there may be in-situ change resprout from underground lignotubers. Species which (Dunlop & Brown 2008), where some species already present resprout may also shed seeds from capsules and recruit in the GBMWHA will be favoured by climate change and seedlings as well. A few species most notably Eucalyptus may increase in abundance, while others may decline. The oreades are killed by fire and release their seeds into the result of this process is likely to be shifts in the relative ground that has been cleared by flame and made more fertile abundance of different eucalypt species. with ash. An interesting aspect of the reproductive strategy of Eucalyptus oreades is early seed production in trees with In the Blue Mountains much of the wet sclerophyll forest suppressed growth relative to dominant trees. This acts as a occurs as relatively small patches within a matrix of drier hedge against another fire in the period before the dominant forest. These patches are vulnerable to regular fire, especially trees reach maturity (Glasby et al, 1988). where their topographic position does not offer protection from fire. The shale cap forests of the Blue Mountains are Some species such as the Angophora species shed their seed especially vulnerable because they occur on hill tops. The wet annually with recruitment depending on rainfall conditions. forests occurring as narrow bands along watercourses in the Indeed rather than being particularly fire-dependent species sandstone gorges are similarly vulnerable. The loss of limbs as is sometimes claimed, many are dependent on disturbance and foliage in dry periods may contribute to fire risk. Some conditions that provide suitable conditions for seedling canopy dominants typical of wet sclerophyll forest are weak recruitment e.g. the creation of bare areas following soil resprouters (e.g. Eucalyptus deanei). Hence, they would be movement, flood alluvium, grazing, drought or sometimes vulnerable to regular, intense fire. Regeneration of Eucalyptus fire. Saplings will often be seen on the edges of disturbed deanei from seed may also be compromised by intense fire. It sites such as roadsides. Eucalyptus oreades will successfully sheds its seed at maturity, with germination mainly occurring establish along roadsides and in gardens in the absence of on bare ground that has been exposed following a low intensity fire (DHB pers obs.). fire or other periodic disturbance. Its capsules are not very woody and would be likely to burn in an intense crown fire. Biogeographic variation On the other hand it is possible that some capsules and seed may be blown long distances in the updraughts associated The GBMWHA extends from the Hunter Valley in the north with such a fire, as are leaves and twigs. to the Southern Highlands and is centred on the Central Coast and Central Tablelands botanical divisions of NSW, but includes parts of the Central Western Slopes and the North Conclusion Coast divisions. The northern part of the region includes some eucalypts such as Eucalyptus laevopinea that are The 96 eucalypts (55 widespread, 41 restricted) known more widely distributed in northern NSW and Queensland. from the GBMWHA occupy a very wide range of habitats A disjunct population of Eucalyptus nobilis, a species more being really only absent from rainforest and swamp sites. typical of the Northern Tablelands, has been discovered near The majority of species have components of both stress- Mount Coricudgy during a current survey of Wollemi NP tolerator and competitor strategies and this has probably (Stephen Bell pers. com.) will not be included in the list until been a main contributor to their success. While we have been the survey and identifications have been completed. In the Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area 443 able to make some generalisations about the habitat of many References of the eucalypts, details of the ecology of most species are still poorly-known. How they will respond to the changed Bell S.A.J. (1998) Wollemi National Park Vegetation Survey. A fire conditions as temperatures increase, available moisture management document. (East Coast Flora Survey Warners Bay changes and fire patterns change is impossible to predict with NSW). our current knowledge. To guide management in conserving Bell, S.A.J. (2001) Notes on the distribution and conservation status all these species for future generations we definitely need of some restricted plant species from sandstone environments more research on the ecology of these dominants of the of the upper Hunter Valley, New South Wales Cunninghamia Australian forests. 7 (1): 77–88. Bell, S.A.J. (2008) Rare or threatened vascular plant species of Wollemi National Park central eastern New South Wales Cunninghamia 10 (3): 331–371. Acknowledgements Bell, S.A.J., Klaphake, V., & Nicolle, D. (in prep) Eucalyptus expressa (Myrtaceae): a distinctive new species from the The vegetation surveys within the GBMWHA were funded sandstone ranges north-west of Sydney, New South Wales. MS by the Central Branch of the Parks & Wildlife Group submitted to Telopea (2010). within DECCW, and by the Sydney Catchment Authority. Benson, D.H. (1985) Aspects of the ecology of a rare tree species We would like to thank Daniel Connolly, Stephen Bell Eucalyptus benthamii at Bents Basin, Wallacia. Cunninghamia 1 (3): 371–383. and Peter Myerscough for their advice and comments on Benson, D.H. (1992) The natural vegetation of the Penrith 1:100 the manuscript. We would also like to thank Bob Conroy 000 map sheet. Cunninghamia 2 (4): 503–662. for initially encouraging us to prepare the definitive list of Benson, D. & McDougall, L. (1998) Ecology of Sydney plants. Part eucalypts, and for his support for the biodiversity survey 6 Dicoyledon family Myrtaceae. Cunninghamia 5 (4): 808–987. program within the GBMWHA. Thanks are also due to the Black, D. (1982) The vegetation of the Boyd Plateau N.S.W. Plant botanists who have undertaken the (often arduous) field Ecology 50 (2): 93–111. surveys. Briggs, J.D. & Leigh, J.H. (1996). Rare or threatened Australian plants (Revised Edn; CSIRO: Australia).

Fig. 5. Unlike most eucalypt species, mature Eucalyptus oreades trees (Blue Mountains Ash) are killed by fire; vigorous post-fire seedling recruitment leading to groves of even-aged saplings as here in the misty forests of Blue Mountains National Park. 444 Cunninghamia 11(4): 2010 Hager & Benson, Eucalypts of the Greater Blue Mountains World Heritage Area

Brooker, M.I.H & Kleinig D.A. (1983) Field Guide to Eucalypts. Peake, T.C. (2006) The vegetation of the Central Hunter valley, Volume 1. South-eastern Australia. Inkata Press, Melbourne. New South Wales. A report on the findings of the Hunter DEC (2004) The native vegetation of the Nattai and Bargo Remnant Vegetation Project. (Hunter-Central Rivers Catchment reserves. Unpublished report. (NSW Dept of Environment & Authority, Paterson). Conservation, Hurstville). http://www.environment.nsw.gov. Pfeil, B.E. & Henwood, M.J. (2004) Multivariate analysis of au/surveys/NattaiNPEtalNativeVegetation.htm morphological variation in Eucalyptus series Psathyroxyla DEC (2006) The vegetation of the western Blue Mountains. Blakely (Myrtaceae): taxonomic implications. Telopea 10(3): Unpublished report funded by the Hawkesbury – Nepean 711–724. Catchment Management Authority. (NSW Dept of Environment PlantNet – the Royal Botanic Gardens Online Flora of New South & Conservation, Hurstville). Wales http://plantnet.rbgsyd.nsw.gov.au/floraonline.htm DEC (2006a) The native vegetation of the Woodford and Erskine Pryor, L.D. & Johnson, L.A.S. (1971) A classification of the Ranges, Kings Tableland and Narrow Neck Peninsula in Blue eucalypts. (Australian National University Press: Canberra). Mountains National Park. (NSW Dept of Environment & Conservation, Hurstville). Manuscript accepted 4 November 2010 DECC (2008) The native vegetation of the Yengo and Parr Reserves and surrounds. (NSW Dept of Environment & Climate Change NSW, Hurstville). Dunlop, M. & Brown, P.R. (2008) Implications of climate change for Australia’s National Reserve System: A preliminary assessment. Report for the Department of Climate Change. (Dept of Climate Change, Canberra). Fairley, A. & Moore P. (2000) Native plants of the Sydney District, an identification guide. (Kangaroo Press, Roseville NSW). Glasby, P., Selkirk, P.M., Adamson, D., Downing, A.J., & D.R Selkirk (1988) Blue Mountains Ash (Eucalyptus oreades R. T. Baker) in the Western Blue Mountains. Proceedings of the Linnaean Society of New South Wales 110 (2): 141–158. Grime J.P. (2001) Plant strategies, vegetation processes and ecosystem properties. 2nd edition. (John Wiley & Sons: Chichester). Hill, K.D. (2002) Angophora; Corymbia; Eucalyptus In Harden, G.J. (Ed.) Flora of New South Wales Volume 2 Revised Edit. (UNSW Press: Sydney). Hill, K.D. & Johnson, L.A.S. (1995) Systematic studies in the eucalypts. 7. A revision of the bloodwoods, genus Corymbia (Myrtaceae). Telopea 6: 185–504. Holland, W.N., Benson, D.H & McRae, R.H.D. (1992) Spatial and temporal variation in a perched headwater valley in the Blue Mountains: geology, geomorphology, vegetation, soils and hydrology. Proceedings of the Linnean Society of NSW 113: 271–295. Johnson, L.A.S. & Hill, K.D. (1990) New taxa and combinations in Eucalyptus and Angophora (Myrtaceae). Telopea 4(1): 37–108 Keith, D.A & Benson, D.H. (1988) The natural vegetation of the Katoomba 1:100 000 map sheet. Cunninghamia 2(1):107–144. McKinnon, G.E., Jordan, G.J., Vaillancourt, R.E., Steane, D.A. & Potts, B.M. (2004) Glacial refugia and reticulate evolution: the case of the Tasmanian eucalypts. Philosophical Transactions of the Royal Society of London 359: 375–284.DOI 10.1098/ rstb.2003.1391 Myerscough, P.J. (1998) Ecology of Myrtaceae with special reference to the Sydney region. Cunninghamia 5 (4): 787–807. NPWS (1998) The Greater Blue Mountains Area World Heritage Nomination (NSW National Parks & Wildlife Service and Environment Australia). NPWS (2003) The native vegetation of the Warragamba Special Area. (National Parks & Wildlife Service, Hurstville and Sydney Catchment Authority, Penrith).