Research Paper |
Corresponding author: John T. Hunter ( jhunter8@bigpond.com ) Academic editor: Jorge Capelo
© 2022 John T. Hunter, Donna Lewis, Eda Addicott, Sarah Luxton, Ian Cowie, Ben Sparrow, Emrys Leitch.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Hunter JT, Lewis D, Addicott E, Luxton S, Cowie I, Sparrow B, Leitch E (2022) A plot-based analysis of the vegetation of the Northern Territory, Australia: a first assessment within the International Vegetation Classification framework. Vegetation Classification and Survey 3: 161-174. https://doi.org/10.3897/VCS.83045
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Aims: To develop an interim classification of the vegetation of the Northern Territory at the International Vegetation Classification (IVC) division (level 4) and macrogroup (level 5) levels. These types are produced to assist in the development of an integrated nationwide plot and floristically based classification of Australia allowing integration within a global perspective. Study Area: The Northern Territory of Australia covers an area of 1.42 million square kilometres, almost 20% of Australia’s land mass. It comprises three distinct climatic zones including tropical, subtropical and arid vegetation types. Methods: We used collated vegetation data held by two organisations: the Northern Territory Government, Department of Environment, Parks and Water Security and the Terrestrial Ecosystem Research Network (a total of 45,710 plots used). We applied semi-supervised quantitative classification methods to define vegetation types at the IVC division and macrogroup levels. Analyses used kR-CLUSTER methods on presence/absence data. Macrogroups were characterised by taxa with the highest frequency of occurrence across plots. Additional analyses were conducted (cluster) to elucidate interrelationships between macrogroups and to assist in the assessment of division level typology. Results: We propose 21 macrogroups and place these within higher thematic levels of the IVC. Conclusions: We found that the IVC hierarchy and associated standard procedures and protocols provide a useful classification tool for Australian ecosystems. The divisions and macrogroups provide a valid framework for subsequent analysis of Northern Territory vegetation types at the detailed levels of the IVC. A consistent typology for the Northern Territory (and hopefully in future, for all of Australia) has numerous benefits, in that they can be used for various applications using a well-structured, systematic and authoritative description and classification that is placed in a continental and global context, readily enabling the one system to be used in studies from the local to global level.
Taxonomic reference:
Abbreviations: DVT = Definitive Vegetation Type; IVC = International Vegetation Classification; nMDS = non-metric multidimensional scaling; NT = Northern Territory; NTVSD = Northern Territory Vegetation Site Database; NVIS = National Vegetation Information System; WA = Western Australia.
arid, Australia, Definitive Vegetation Type, International Vegetation Classification, National Vegetation Information System, Northern Territory, semi-arid, sub-tropical, tropical
A unified terminology and procedure for classifying vegetation types across jurisdictions and continents is required for clear understandings of their distribution, evolutionary biology, and threats, along with guiding their restoration and rehabilitation (
Australia’s vegetation is unique with more than 80% of the vascular flora endemic to the continent (
Although no continental plot-based vegetation classification currently exists for classifying community types, the Australian National Vegetation Information System (NVIS) is an existing national classification. It is a supervised, largely structural, mapping-based classification with limited floristic data. NVIS was established by the Executive Steering Committee for Australian Vegetation Information (
The NT is a semi-independent jurisdiction covering 20% of the Australian continent. The mid-1980s saw the introduction of vegetation mapping reliant on plot data to classify vegetation types into mappable units. Ad hoc vegetation mapping continues in the NT at varying degrees of attribute detail and spatial scale. No strategic vegetation mapping program for the whole of the NT has been implemented, although scoping documents have been devised to map the NT at 1:100,000 on several occasions (
A more recent attempt to classify the vegetation communities of the NT was conducted in 2007 (
Although initial plot-based data collection within the NT began in the 1950s, the greatest efforts were placed during the early 2000s; however, no centralised database was available for the collation of early data (
The IVC is based on the EcoVeg approach (
The IVC has been developed with a rigorous set of standards and principles that seek to characterise the world’s vegetation. To date, the IVC has been built to its greatest depth in North America, including national vegetation classifications (NVCs) in both the United States (USNVC) and Canada (
Here we use semi-supervised analyses incorporating plot data from across the NT in order to circumscribe a preliminary series of mid-level (L4 division and L5 macrogroup) IVC natural vegetation types. These types are placed where possible within existing and proposed IVC types, such as those published in
The NT of Australia comprises an area of 1.42 million square kilometres, one-sixth of the total land area of Australia and is sparsely populated (244,800 people; 60% living within the 3,164 square kilometres of the Greater Darwin region (the capital);
Locality map of Australia’s Northern Territory, illustrating the distribution of 2,245 vegetation plots to inform the 1990 1:1 million vegetation map and the additional 45,710 plots sampled in this analysis. Bioregions from interim Biogeographic Regionalisation for Australia (IBRA) version 7 (2012). NTVSD – Northern Territory Vegetation Site Database; TERN – Australian Terrestrial Ecosystem Research Network.
The Interim Biogeographic Regionalisation for Australia (IBRA) provides a division of Australia into units of broadly similar landform, geology and biodiversity akin to ecoregions. These ‘bioregions’ provide a useful unit for natural resource management and planning (
The NT is characterised by tropical and arid zone vegetation types. Tropical savannas are the dominant vegetation type of northern Australia covering about 25% of the continent (
The data extracted from the NTVSD used a common survey protocol. The standard method for vegetation sampling in the NT has been consistently used by government botanists and scientists since the 1970s (
The Terrestrial Ecosystem Research Network (TERN) Ecosystem Surveillance monitoring plots of 100 m × 100 m were also used in the analyses. A sampling plot includes the same floristic, structural and environmental attributes as that collected on the NTVSD, however the methods differ and are more rigorous for monitoring purposes as outlined in
The full set of more than 77,000 plots were accessed from the NTVSD 2021. The plot data is of varied quality and detail with information including full floristic or dominant species only and data scored on presence/absence or percent cover (
Datasets without full floristic information or of plot sizes less than 100 square metres or greater than 25,000 square metres were removed from the NTVSD dataset. Plots where species richness was less than 10 taxa were also removed. It is acknowledged that the removal of plots with less than 10 taxa is likely to remove from the analysis some distinct, but low diversity communities such as coastal samphire, mangrove and some freshwater floodplain vegetation. However, our analysis is not designed to develop a definitive set of all possible types through analysis, but to provide an initial set of types which will be expanded with further analysis both within the NT and across jurisdictional boundaries.
Records that were not identified to species level or better were excluded as they can artificially inflate species richness, as can plots with low frequency taxa. As the remaining dataset was a mixture of plots where taxa were recorded by percentage cover or presence/absence, all scores were reduced to presence/absence only. A total of 45,710 plots were used in subsequent analyses representing 4,566 taxa and 831,149 records. Taxonomic curation was applied to the vascular plants presence data to ensure names to the species-rank were current and in accordance with the Australian Plant Census (APC: Council of Heads of Australasian Herbaria (CHAH)), various dates) and the HOLTZE taxon table, the authoritative vascular plant checklist for the NT as available through Flora NT (
Our analysis utilises a broad concept of Coolabah – Eucalyptus microtheca, which here includes a number of segregate taxa recognised by
Growth form data, in accordance with the NVIS growth form definitions and controlled vocabulary, were updated and extracted from HOLTZE for each species (
Due to size of the final dataset and the computation power required, a limited set of options were available to the authors for initial analyses. Data were transformed into a species by site matrix via the r-package ‘tidyr’ (v 1.1.3;
A combination of statistical and intuitive approaches can often produce accurate and stable outcomes particularly at the mid to higher hierarchical levels of a classification (
An important component of this work was to align our results with the IVC hierarchy. To do this we used the key to IVC formation classes and brief definitions provided by
Four broad types were recognised from initial analysis and can be compartmentalised into the NTs climatic zones (based on Koppen-Geiger:
Floristic relatedness of macrogroups based on the 150 most frequent taxa across all plots used to define each type. Only macrogroups defined within this current investigation appear in the analysis. Note this analysis was used as a guide for placement within the upper IVC hierarchy, but due to necessary incorporation of growth form data at the macrogroup and higher levels, these results do not directly relate to the final placement. See Table
Due to the necessary inclusion of growth form and climate the final alignment with the IVC hierarchy does not follow the results of the floristic relatedness as indicated in Fig.
Alignment of the Northern Territory vegetation types with the International Vegetation Classification hierarchy existing Class, Subclass, Formation, and proposed preliminary Divisions and Macrogroups.
Class | Subclass | Formation | Division | Macrogroup | Analysis Group |
---|---|---|---|---|---|
1. Forest & Woodland | |||||
1.A. Tropical Forest & Woodland | |||||
1.A.1. Tropical Dry Forest & Woodland | |||||
L.4. Australasian Dry Forest & Woodland NEW | |||||
L.5 (MG1) Australasian & East Malesian Dry Forest NEW | [20-18] | ||||
1.A.5 Mangrove | |||||
1.A.5.Wb Indo-West Pacific Mangrove | |||||
L.5 (MG2) West Pacific (East Melanesia, Micronesia, Polynesia) Mangrove (M208) | |||||
2. Shrub & Herb Vegetation | |||||
2.A. Tropical Grassland, Savanna & Shrubland | |||||
2.A.1. Tropical Lowland Grassland, Savanna & Shrubland | |||||
L.4 Australian Tropical Savanna and Scleromorphic Woodland (D133; |
|||||
L.5 (MG3) Australian Paperbark Melaleuca viridiflora–Long-fruited Bloodwood Corymbia polycarpa Forest and Woodland NEW | [20-2] | ||||
L.5 (MG4) Australian Darwin Stringybark Eucalyptus tetrodonta Scleromorphic Woodland (M530; |
[20-6,14,15,19] | ||||
L.5 (MG5) Australian Darwin Box Eucalyptus tectifica Scleromorphic Woodland NEW | [20-4] | ||||
L.5 (MG6) Australian Broad-leaved Bloodwood Corymbia foelscheana Scleromorphic Woodland NEW | [20-3] | ||||
L.5 (MG7) Australian Lancewood Acacia shirleyi Forest, Woodland and Shrubland NEW | [20-5] | ||||
L.5 (MG8) Australian Coolabah Eucalyptus microtheca Tropical Savanna Grassy Woodland and Tussock Grassland NEW | [20-7] | ||||
L.5 (MG9) Australian Tropical Triodia bitextura–Triodia microstachya–Triodia bynoei Hummock Grassland and Open Woodland (M531; |
|||||
L.5 (MG10) Australian Small-fruited Bloodwood Corymbia dichromophloia–Curley Spinifex Triodia bitextura Scleromorphic Woodland NEW | [20-13] | ||||
L.5 (MG11) Australian Tropical Billy Goat Plum Terminalia carpentariae–Northern Spinifex Triodia microstachya Scleromorphic Woodland and Shrubland complex NEW | [20-20] | ||||
L.5 (MG12) Australian Bean Tree Bauhinia cunninghamii Tropical Savanna Tussock Grassy Woodland and Tussock Grassland NEW | [20-1,8,16] | ||||
L.5 (MG13) Eremaean Semi-arid Hummock Grassland and Low Open Woodland (M532; |
|||||
2.C.3 Tropical Freshwater Marsh, Wet Meadow & Shrubland | |||||
L.4 Australian Tropical Grassland and Sedgeland NEW | |||||
L.5 (MG14) Northern Australia Tropical Swamp Grass Pseudoraphis spinescens–Water Chestnut Eleocharis dulcis–Water Lily Nymphaea violacea Grassland and Sedgeland NEW | [20-9] | ||||
2.C.4 Temperate to Polar Freshwater Marsh, Wet Meadow & Shrubland | |||||
L.4 Australian Arid and Semi-arid Grasslands and Sedgelands NEW | |||||
L.5 (MG15) Australian Ephemeral Arid and Semi-arid wetlands NEW | Arid Wetlands | ||||
3. Desert & Semi-Desert | |||||
3.A. Warm Desert & Semi-Desert, Scrub & Grassland | |||||
3.A.2. Warm Desert & Semi-Desert, Scrub & Grassland | |||||
L.4 Australian semi-desert scrub and grassland (D330; |
|||||
L.5 (MG16) Australian Desert Hummock Grassland (M535; |
|||||
L.5 (MG17) Australian Desert and Semi-arid Mulga Acacia aneura Woodland & Shrubland NEW | [10-10] | ||||
L.5 (MG18) Australian Desert and Semi-arid Victoria Wattle Acacia victoriae–Emu Bush Eremophila longifolia–Ironwood Acacia estrophiolata Shrubland NEW | [10-1] | ||||
L.5 (MG19) Australian Desert and Semi-arid Witchetty Bush Acacia kempeana Shrubland NEW | [10-5] | ||||
L.5 (MG20) Australian Desert and Semi-arid Bloodwood Corymbia opaca Woodland NEW | [10-8] | ||||
L.5 (MG21) Australian Desert and Semi-arid Rock Fuchsia Bush Eremophila freelingii– Witchetty Bush Acacia kempeana Shrubland NEW | [10-7] | ||||
5. Aquatic Vegetation | |||||
5.B Freshwater Aquatic Vegetation | |||||
5.B.1 Tropical Freshwater Aquatic Vegetation | |||||
L.4 Australian Tropical Aquatic Vegetation NEW | |||||
L.4 Australian Desert and Semi-arid Aquatic Vegetation NEW |
The NT vegetation types were aligned within the three highest levels of the IVC hierarchy, which are global in scope (formation class, formation subclass and formation). At the highest level, the NT vegetation types could be placed into the Formation Classes 1. Forest and Woodland, 2. Shrub & Herb Vegetation, 3. Desert & Semi-Desert and 5. Aquatic Vegetation. The majority of the types (14 of 21) occur in the tropical and subtropical climatic zones, which cover approximately half of the NT. Within these zones, 11 of the 14 types were placed as macrogroups into: formation class 2. Shrub and Herb Vegetation, formation subclass 2.A. Tropical Grassland, Savanna and Shrubland, formation 2.A.1 Tropical Lowland Grassland, Savanna & Shrubland, and into a specific division for Australian tropical savannas: L.4 Australian Tropical Savanna and Scleromorphic Woodland, as defined by
Analysis group MG1, broadly categorised as ‘closed forest’, was the only vegetation type placed into the IVC formation class 1. Forest & Woodland, formation subclass 1.A. Tropical Forest & Woodland, and formation 1.A.1. Tropical Dry Forest & Woodland (Table
Further south in the arid zone, six vegetation types were placed into formation class 3. Desert & Semi-Desert, formation subclass 3.A. Warm Desert & Semi-Desert, Scrub & Grassland and formation 3.A.2. Warm Desert & Semi-Desert, Scrub & Grassland. A recent study by
Suppl. material
Here we present the classification of a large vegetation plot dataset (45,710 plots) from the NT, Australia, and align results with the upper levels of the IVC (macrogroup to formation class). Half of the vegetation types (11 of 21 macrogourps) fall into the formation 2.A.1 Tropical Lowland Grassland, Savanna & Shrubland and an Australian specific division – L.4 Australian Tropical Savanna and Scleromorphic Woodland. Closed Forest sites were the only type to align with formation class 1. Forest & Woodland, formation subclass 1.A. Tropical Forest & Woodland, with the potential for a new division L.4. Australasian Dry Forest & Woodland and macrogroup L.5 (MG1) Australasian & East Malesian Dry Forest; that reflects the areas affiliation with the South East Asian flora. Various types covering smaller areas were identified, including one type in formation 2.C.3 Tropical Freshwater Marsh, Wet Meadow & Shrubland, which is dominated by tropical grasses and sedges. Further south, arid types, including woodlands, shrublands and grasslands aligned well with the proposed division L.4 Australian semi-desert scrub and grassland (D330). While NT-derived data underpins this study types within the tropical, subtropical and arid regions cross into WA, Qld, SA, and NSW, highlighting the potential of the IVC as a cross-jurisdictional framework for a nationally consistent vegetation classification in Australia.
One of the strengths of the IVC is that it provides a framework for both existing types and ‘place holders’ to enable debate on the appropriateness of a type in the hierarchy. For example, the distinctive mangrove vegetation can be aligned with 1. A.5.Wb Indo-West Pacific Mangrove division and the L.5 (MG2) West Pacific (East Melanesia, Micronesia, Polynesia) Mangrove (M208) macrogroup, based on
The assignment of new types to the IVC was recently demonstrated by
In total, of the eight divisions and 21 macrogroups described across the NT, five new divisions and 16 macrogroups are proposed. The need for these additional groups reflects the high endemicity of the Australian flora (80% of taxa are endemic) and largely northern hemisphere focus of the IVC to date. Of the proposed macrogroups, NT botanical experts suggest that East Malesian (and Australian) Eucalyptus alba savanna likely fits under one of the macrogroup L5 categories. However, the ground layer may be too similar to other eucalypt communities for it to be differentiated and its area of occupancy is small. Additionally, there is significant overlap in the East Malesian – north Australian grasses and more plot data would be needed to substantiate these inferences. In addition to the IVC’s flexibility, which enables the incorporation of both existing and new types, it provides a numerical, plot-based framework for defining lower-level types in the hierarchy. This is important in the NT, where much historic work has been done, but modern analysis tools have not been widely implemented.
An additional usage for the IVC in the NT is as a framework to bridge historic work with modern vegetation science. Existing classification systems include the 1:1,000,000 vegetation map (
The IVC provides a means to undertake cross-jurisdictional vegetation analysis in Australia. The NT occupies 20% of the continent and borders most other major states including WA, Qld, NSW, and SA. A number of our vegetation types span political borders, including the tropical savanna biome (
This analysis highlights how a rigorous rule-based hierarchical classification system, where the lower schematic levels are based on plot-based vegetation analyses of floristic and ecological data, is best placed to underpin our understanding of Australian vegetation. Such processes allow for a better understanding of distribution, interrelatedness and rarity. Lack of clear guidelines and dissimilar processes applied across state and territory borders only adds further confusion leaving practitioners to rely on intuition and opinion. Using a classification system such as the IVC provides an understanding of vegetation types both at local and regional levels and within a continental and global perspective. Our analysis has shown clearly that the IVC can be applied to a significant proportion of the Australian continent at the macrogroup level.
Northern Territory Vegetation Site Database (NTVSD), http://www.ntlis.nt.gov.au/vsd/f?p=113
Natural Resource (NR) Maps data visualisation tool > vegetation > vegetation sites https://nrmaps.nt.gov.au/nrmaps.html
Bookmark NTVSD https://nrmaps.nt.gov.au/nrmaps.html#11569ec5-9ed5-46ce-a084-59bfd049fadc (created December, 2021)
Terrestrial Ecosystem Research Network (TERN) EcoPlots https://ecoplots.tern.org.au
Images of Northern Territory macrogroups https://drive.google.com/drive/folders/1Bl97PYS194XwLZT6c9Kcd-a0HrprJNxH?usp=sharing
Initial data cleaning was provided by DL, subsequent data management and analyses were conducted by JH. JH and DL defined types and their placement within the IVC hierarchy. All authors were involved equally in the assessment of type validity, final circumscription, naming and in the writing of the manuscript.
The authors are indebted to Don Faber-Langendoen from NatureServe for providing guidance in aligning NT vegetation types to the IVC hierarchy at the mid-levels and assisting with nomenclature for the new divisions and macrogroups. Various observers from the NT Government who collected vegetation plot-based data from across the NT are acknowledged. Particular thanks are extended to NT Herbarium staff (past and present) for ongoing curation and maintenance of the NT flora and vegetation datasets, including taxonomists, botanists and other scientists. The authors are grateful to NT flora and vegetation experts; Catherine Nano, Peter Jobson, Nick Cuff, Diane Napier and Dominique Lynch for their review, as a result, the new NT divisions and macrogroups are considered more robust. The TERN cohort are thanked wholeheartedly for providing data, representative photographs, and future Australian standards for vegetation plot-based data especially Sally O’Neill and Siddeswara Guru.
Classification, diagnostic taxa and notes on the distribution of Northern Territory Macrogroups and synoptic table.