New paper: Long-term patterns of invertebrate abundance and relationships to environmental predictors factors in arid Australia

Authors: Alan B.C. Kwok, Glenda M. Wardle, Aaron C. Greenville, Chris R. Dickman.

Published in: Austral Ecology

This paper represents the first published study from the Desert Ecology Research Group on the invertebrates that occur in our study region in the Simpson Desert. Even though we have been surveying invertebrates for over two decades for various projects, Alan was able to collate all the data and get it (and us) organsied!



Resource pulses are a key feature of semi-arid and arid ecosystems, and are generally triggered by rainfall. While rainfall is an acknowledged driver of the abundance and distribution of larger animals, little is known about how invertebrate communities respond to rain events or to vegetative productivity. Here we investigate Ordinal-level patterns and drivers of ground-dwelling invertebrate abundance across six years of sampling in the Simpson Desert, central Australia. Between February 1999 and February 2005, a total of 174,381 invertebrates were sampled from 32 Orders. Ants were the most abundant taxon, comprising 83% of all invertebrates captured, while Collembola at 10.3% of total captures, were a distant second over this period. Temporal patterns of the six invertebrate taxa specifically analysed (Acarina, ants, Araneae, Coleoptera, Collembola and Thysanura) were dynamic over the sampling period, and patterns of abundance were taxon-specific. Analyses indicate that all six taxa showed a positive relationship with the cover of non-Triodia vegetation. Other indicators of vegetative productivity (seeding, flowering) also showed positive relationships with certain taxa. Although the influence of rainfall was taxon-dependent, no taxon was affected by short-term rainfall (up to 18 days prior to survey). The abundance of Acarina, ants, and Coleoptera increased with greater long-term rainfall (up to 18 months prior to survey), whilst Araneae showed the opposite effect. Temperature and dune zone (dune crest vs. swale) also had taxon-specific effects. These results show that invertebrates in arid ecosystems are influenced by a variety of abiotic factors, at multiple scales, and that responses to rainfall are not as strong or as predictable as those seen for other taxa. Our results highlight the diversity of invertebrates in our study region, and emphasize the need for targeted long-term sampling to enhance our understanding of the ecology of these taxa and the role they play in arid ecosystems.


Kwok, A. B. C., G. M. Wardle, A. C. Greenville, and C. R. Dickman. (2016). Long-term patterns of invertebrate abundance and relationships to environmental factors in arid Australia. Austral Ecology 41: 480-491.

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New paper: Cattle removal in arid Australia benefits kangaroos in high quality habitat but does not affect camels

Authors: Anke S. K. Frank, Glenda M. Wardle, Aaron C. Greenville and Chris R. Dickman

Published in: The Rangeland Journal


Removing cattle as a management tool to conserve biodiversity may not necessarily alter grazing impacts on vegetation if other introduced or native herbivores move in and replace the cattle after removal. This study investigated whether there was a difference in the abundance of native red kangaroos (Osphranter (Macropus) rufus) and introduced feral camels (Camelus dromedarius) on arid rangelands where cattle had been recently removed compared with where cattle remained. Activity was measured by clearing and weighing dung, and by counting animal sightings. Kangaroos were encountered more frequently in high quality habitat (gidgee woodland) where cattle had been recently removed. However, kangaroo dung in newly cattle-free areas comprised only ~1.5% of the weight of cattle dung in this habitat where cattle still grazed, indicating no grazing compensation by the native herbivore. Camels showed no clear preference for particular habitat types but used dune tops usually avoided by kangaroos and cattle. There was no indication of camels using habitats differently in areas where cattle were removed. Camel dung collected across all habitats comprised less than a tenth the weight of cattle dung, but almost five times as much as kangaroo dung. As cattle removal had occurred relatively recently, further monitoring is needed to determine its impact over longer periods,  especially through low and high rainfall cycles. Methods to improve the monitoring of large herbivores in the presence and absence of livestock and to assess whether anticipated conservation goals are achieved are discussed.achieved.


Frank A. S. K., Wardle G. M., Greenville A. C. & Dickman C. R. (2016). Cattle removal in arid Australia benefits kangaroos in high quality habitat but does not affect camels. The Rangeland Journal 38: 73-84.


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2014 ‘Ecology in Action’ Photographer of the Year

The Ecological Society of Australia “Ecology in Action” Photographic Competition is an annual event attracting amazing images celebrating the diversity of landscapes and ecosystems within Australia and New Zealand. In 2014, a new award was added for Best Portfolio and attracted a cash prize as well as the opportunity for an online photographic exhibition of the winner’s work. The inaugural prize was shared by Dr Aaron Greenville from the University of Sydney and Richard Wylie, based at Monash University, Victoria.

Head to the society’s webpage to see my online exhibition featuring the images which won me best portfolio in the 2014 competition, as well as some additional images showcasing  my photographic interests.

A mulgara from the Simpson Desert, Qld, Australia.

A mulgara from the Simpson Desert, Qld, Australia.


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ESA 2015 conference talk: Managing species across vast spatial areas: does one size fit all?

Understanding how the spatial and temporal dynamics of populations vary across the landscape is fundamentally important to managing and conserving species. For example, populations may fluctuate in synchrony, or exhibit other forms of spatial sub-structuring, due to intrinsic population parameters or to influences from environmental factors. Importantly, synchronous populations may be at greater risk of extinction if all populations are decreasing to zero at the same time, thereby reducing rescue through colonisation. Different species may not have similar dynamics, even if they share the same environment and thus unravelling the spatial dynamics of multiple species provides vital information about what scale to apply management actions.

MARSS models
MARSS framework is hierarchical and allows modelling of different spatial population structures and parameters, such as density dependence, while including both process and observation variability. Process variability represents temporal variability in population size due to environmental stochasticity. Observation variability includes sampling error. The process component is a multivariate first-order autoregressive process and is written in log-space:
MARSS process eqnwhere X = matrix of all m sub-populations at time t
B = density dependence
u = mean growth rate of the sub-population
w = process errors, assumed to be independent and to follow a multivariate normal distribution with a mean of 0 and variance-covariance matrix Q.
The observation component, written in log-space:
MARSS obs eqnwhere Y = a matrix of observations of all sub-populations at time t,
a = the mean bias between sites
Z = a matrix of 0’s and 1’s that assigns observations to a sub-population structure.
v = observation error, assumed to be uncorrelated and follow a multivariate normal. distribution, with a mean of 0 and a variance-covariance matrix R

Using long-term data (17‑22 years) across a large-scale study region (8000 km2) in arid central Australia, we test for regional synchrony in a population driver, annual rainfall, across nine sites (>20 km apart). We then draw from examples from small mammal and reptile populations and investigate if each species exhibits synchrony. For species that did not exhibit synchrony, we used multivariate autoregressive state-space (MARSS) models to explore four other sub-population structures. We also use the MARSS models to identify important drivers that may regulate populations of these species.

We show that species exhibited different spatial population structuring and respond to extrinsic factors in different ways. We conclude that investigating how the spatial connections among populations interact with their temporal dynamics and eventual persistence or decline, is important for determining the appropriate scale to implement management actions and that “one size does not fit all”.

More on population dynamics of small mammals, MARSS models and Moran:

EcoTas 2013: Spatial and temporal synchrony in small mammal populations


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My PhD journey comes to an end: the role of ecological interactions

Ecology is often described as the web of life. Many complex interactions occur across this web, including between individuals of the same or different species (biotic) or between the environment and a species (abiotic). Understanding ecological interactions, such as the role of predators in an ecosystem or how climate influences the abundance of a species, are fundamental for describing how this web of life works.

After working for over 10 years in central Australia I had become hooked on the Simpson Desert. The sand dunes of the Simpson Desert that lie in south-western Queensland and towards the border of the Northern Territory are like a second home to me. The red parallel dunes roll across the landscape like waves across the ocean, carrying spinifex, a spiky grass that form odd-looking donuts floating the red surface. There is something about the red sand that gets under your skin and keeps drawing you back. This setting was where I would tackle the question—what are the relative roles of both biotic and abiotic interactions for driving an ecosystem? The Simpson Desert, like many areas of Australia, has another characteristic I could take advantage of: every trip to the Simpson is different. A small storm may have gently swept rain across the sand and marked out a patch of green for all to see against a canvas of red. Further down the road, the vegetation may lay twisted and grey, prostrate as if it collapsed from the effort of begging for rain. This high variability across time and the landscape must be important for explaining how species live in such a hostile environment. Furthermore, the earth’s climate is changing. How will climate change affect the ecology of central Australia, a region of Australia where the extinctions of native animals has already hit the hardest? What lessons can this remote and understudied part of the globe teach us about our world? It was from these questions pressing on my mind that my PhD was born.


Small storms can fall through-out the year in the Simpson Desert turning patches green, but other areas remain dry. This variability added an extra dimension to my PhD project. Photo by Aaron Greenville.

I found that the operation of both biotic (species‑species) and abiotic (species‑environment) interactions were important for the ecology of the Simpson Desert (see figure below). More interestingly, interactions are not fixed and can change across the landscape and through time. For example, the relative influence of predation compared to increases in food from large rainfall events varied over the years. How this affected the role of the dingo in supressing smaller predators, such as the red fox and feral cat, and the implications for feral predator management were highlighted in an earlier article.

Chapter 8-fig1

Figure 8.1 from my thesis: Summary of thesis findings on biotic and abiotic interactions in an arid environment. Here we see how interactions are context-specific and can change over the landscape and time. Grey arrows are possible future interactions to explore. Diagrams by A. Foster.

Fierce debates raged in 1950s and 1960s the about the importance of biotic verses abiotic factors for populations after some Australian ecologists argued that the established dogma of the northern hemisphere ecologists may not be the whole story. The upstarts, H.G. Andrewartha and L.C. Birch, published a seminal text: The distribution and abundance of animals, which used empirical evidence, rather than the largely theoretical evidence from the northern hemisphere, to demonstrate that the environment (abiotic factors) could influence the population abundance of a species. Today, findings such as mine are showing that both biotic and abiotic factors are important. This kind of understanding is crucial if we are going to try and predict how species will be affected by climate change.

Over the past 100 years the climate of central Australia has changed. The deserts have warmed and the frequency of extreme rainfall events have increased. This may effect populations of native species. For example, extreme rainfall events (>95th quantile, or >400 mm, for the Simpson Desert) were required for rodent populations to increase, resulting in a boom in population numbers. One example was long-haired rat which I wrote about here . However, not all small mammals responded to the same drivers, as dasyurid marsupials (such as dunnarts) were not influenced by extreme rainfall events. These interactions are becoming increasing important to understand, as extreme weather events, such as tropical cyclones, heat waves and flooding rains, are increasing in magnitude and frequency. Increases in extreme weather events may change species’ populations through increases in mortality rates, decreases in reproduction, and by facilitation of invasive species or novel interactions. In addition, extreme weather events, such as flooding rains, are important not only for driving food pulses in arid systems, but also other abiotic events such as wildfire.

Not only do different species show a different response to rainfall over time, they also show different responses across the landscape. Native rodents, like the spinifex hopping mouse, exhibit a high level of population synchrony across the study region. Even though each population was separated by at least 20 km, thus preventing dispersal between populations from being a big factor (the rodents I was studying are only 10‑35 grams and the reptiles are even smaller), large rainfall events that occurred through-out the region could drive all the rodent populations up in the same way. The highs and lows of each population were in synchrony, dancing in uniform across an 8000 km2 study area. The dunnarts did what dunnarts do best: whatever they want. Each population I looked at was dancing to its own beat. Contrasting strategies in responses to rainfall in a water-limited environment. Such information is vital for knowing how best to conserve each species. A “one size fits all” approach to conservation management will not work for the small mammals and reptiles I studied.

Species do not only respond to biotic and abiotic factors differently over the landscape or across time. We also need to look at multiple factors, such as rainfall and wildfire, simultaneously when trying to predict how species will interact with each other and their environment. Projecting my findings that the frequency of extreme rainfall events had increased into the future (the next 100 years), populations of rodents did not increase as much as expected, as increases in wildfire frequency and current levels of introduced predators limited rodent populations. After removing introduced predators and keeping dingoes in the system, rodents could take advantage of increases in extreme rainfall events.


The final product after 3.5 years of work: a thesis and several scientific and popular science articles.

I am truly indebted to all the help and support I have had in this journey. Particularly to my supervisors Professors Glenda Wardle and Chris Dickman, my wife, friends and family. Funding was provided by the Australian Research Council, an Australian Postgraduate Award and the Australian Government’s Terrestrial Ecosystems Research Network. Perhaps it is best to end this post with the last paragraph of my thesis:

“This thesis has attempted to take advantage of over 20 years of ecological research focused on the ecology of central Australia. Of this history, I have been fortunate enough to have been part of the last 15 years of the endeavour, working in various capacities. Even though the hand of extinction has been brought down most strongly in Australia’s arid regions, studies such as this elucidate the complexity of the current ecology of central Australia and the sense of wonder that this environment conveys. This wonder feeds curiosity and surely curiosity, the most noble of human traits, needs to be conserved.”


Further reading:

Andrewartha, H. G. and L. C. Birch. 1954. The distribution and abundance of animals. University of Chicago Press, Chicago, USA.

Dickman , C., G. M. Wardle, J. Foulkes, and N. de Preu. 2014. Desert complex environments, Pages 379-438 in D. Lindenmayer, E. Burns, N. Y. Thurgate, and A. Lowe, editors. Biodiversity and environmental change: monitoring, challenges and direction. CSIRO Publishing, Vic.

Greenville, A. C. 2015. The role of ecological interactions: how intrinsic and extrinsic factors shape the spatio-temporal dynamics of populations. PhD Thesis. University of Sydney, Sydney.

Greenville AC, Wardle GM, Tamayo B, Dickman CR (2014). Bottom-up and top-down processes interact to modify intraguild interactions in resource-pulse environments. Oecologia, 1-10.

Greenville, A.C., Wardle, G.M. and Dickman, C.R. (2013). Extreme rainfall events predict irruptions of rat plagues in central Australia. Austral Ecology, 38: 754–764.

Greenville A. C., Wardle G. M., Dickman Christopher R. (2012). Extreme climatic events drive mammal irruptions: regression analysis of 100-year trends in desert rainfall and temperature. Ecology and Evolution, 2, 2645-2658.

Greenville A. C., Dickman C. R., Wardle G. M. & Letnic M. (2009). The fire history of an arid grassland: the influence of antecedent rainfall and ENSO. International Journal of Wildland Fire, 18, 631-639.

Top Dog: How Dingoes Save Native Animals. Australasian Science, November 2014.


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New paper: Ecosystem risk assessment of Georgina gidgee woodlands

Authors: Glenda Wardle, Aaron Greenville, Anke Frank, Max Tischler, Nathan Emery and Christopher R Dickman.

Published in: Austral Ecology Special Issue: Ecosystem Risk Assessment


Ecosystems across the world, and the biodiversity they support, are experiencing increasing anthropogenic pressure, and many will not persist without intervention. Given their complexity, the International Union for Conservation of Nature has adopted an international standard for ecosystem risk assessment that builds on the strengths of the species-based Red List criteria.

A Gidgee stands against a smoky sunset. Photo by Aaron Greenville

A Gidgee (Acacia georginae) stands against a smoky sunset, Simpson Desert, Qld. Photo by Aaron Greenville


We applied this protocol to the relatively understudied Georgina gidgee woodland ecosystem, which has a patchy but widespread distribution in central Australia. To address the extensive knowledge gaps, we gathered data to provide the first description of the characteristic biota, distribution of dominant species and the processes that support the ecosystem. Criteria evaluated include historical, current and future declines in spatial distribution, the extent and area of occupancy, and disruptions to abiotic and biotic processes. Future declines in suitable habitat were based on key climatic variables of rainfall, temperature and soil substrate. We also quantified the uncertainty in bioclimatic models and scenarios as part of predicting degradation of the abiotic environment.

Overall, we assessed the risk status of Georgina gidgee woodlands as vulnerable based on the degradation of abiotic and biotic processes. Bioclimatically suitable habitat was predicted to decline by at least 30% in eight scenarios over the period 2000 to 2050. Predicted declines in overall suitable habitat varied substantially across all scenarios (7–95%). Pressures from grazing, weed encroachment and altered fire regimes further threaten the ecosystem; therefore, vulnerable status was also recorded for future declines based on altered biotic processes. Accurate mapping and monitoring of the study ecosystem should receive priority to inform conservation decisions, and sustainable grazing practices encouraged. Our findings focus attention on other patchily distributed ecosystems that may also have escaped attention despite their contribution to supporting
unique biodiversity and ecosystem services. It is timely that environmental monitoring  and policy account for these natural assets.

Georgina gidgee woodlands snake through the swales in the Simpson Desert. Photo by Aaron Greenville.

Georgina gidgee woodlands snake through the dune swales in the Simpson Desert. Photo by Aaron Greenville.



Wardle, G. M., Greenville A. C., Frank A. S. K. , Tischer M., Emery N. J. & Dickman C. R. (2015). Ecosystem risk assessment of Georgina gidgee woodlands in central Australia. Austral Ecology 40: 444-459.

Related reading:

Dickman C. R., Greenville A. C., Tamayo B. & Wardle G. M. (2011). Spatial dynamics of small mammals in central Australian desert habitats: the role of drought refugia. Journal of Mammalogy 92, 1193-209.

Frank A. S. K., Wardle G. M. , Dickman C. R. & Greenville AC (2014). Habitat- and rainfall-dependent biodiversity responses to cattle removal in an arid woodland-grassland environment. Ecological Applications, 24:2013–2028.


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New paper: On the validity of visual cover estimates for time series analyses

Authors: Vuong Nguyen, Aaron Greenville, Chris Dickman and Glenda Wardle.

Journal: Plant Ecology

Spinifex (Triodia basedowii) in the Simpson Desert, Qld. Photo by Aaron Greenville.

Spinifex (Triodia basedowii) in the Simpson Desert, Qld. Photo by Aaron Greenville.


Changes in vegetation cover are strongly linked to important ecological and environmental drivers such as fire, herbivory, temperature, water availability and altered land use. Reliable means of estimating vegetation cover are therefore essential for detecting and effectively managing ecosystem changes, and visual estimation methods are often used to achieve this. However, the repeatability and reliability of such monitoring is uncertain due to biases and errors in the measurements collected by observers. Here, we use two primary long-term monitoring datasets on spinifex grasslands, each established with different motivations and methods of data collection, to assess the validity of visual estimates in detecting meaningful trends.

The first dataset is characterised by high spatial and temporal coverage but has limited detail and resolution, while the second is characterised by more intensive sampling but at fewer sites and over a shorter time. Using multivariate auto-regressive state-space models, we assess consistency between these datasets to analyse long-term temporal and spatial trends in spinifex cover whilst accounting for observation error. The relative sizes of these observation errors generally outweighed process, or non-observational errors, which included environmental stochasticity. Despite this, trends in the spatial dynamics of spinifex cover were consistent between the two datasets, with population dynamics being driven primarily by time since last fire rather than spatial location. Models based on our datasets also showed clear and consistent population traces.

We conclude that visual cover estimates, in spite of their potential uncertainty, can be reliable provided that observation errors are accounted for.


Nguyen, V., Greenville A., Dickman C., and Wardle G. 2015. On the validity of visual cover estimates for time series analyses: a case study of hummock grasslands. Plant Ecology, 1-14.


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