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FERAL MAMMALS IN AUSTRALIA’S RANGELANDS: FUTURE THREAT, MONITORING AND MANAGEMENT G. P. Edwards, A. Pople, P. Caley and K. Saalfeld Abstract In this paper we provide an initial brief overview of past changes in the biodiversity of Australia’s rangelands. Following on, we focus on current and future threats to biodiversity posed by feral mammals (predators and herbivores) inhabiting the rangelands, exploring trends in populations and options for management. Notably, rabbits have declined in recent years in the wake of Rabbit Haemorrhagic Disease, populations of camels have increased dramatically, while foxes appear to have moved northwards thereby threatening native fauna within their expanded range. Finally we examine how to monitor the impacts of feral mammals so that management can be applied at the correct time and scale. Factors which need to be considered when designing a monitoring programme are discussed. While it is pest impact that should ideally be monitored, this is rarely achieved in practice. Rather, monitoring usually involves population assessments, the untested assumption being that higher densities equate to higher impacts. Current ‘best practice’ methods of monitoring populations of feral mammals in the rangelands are discussed briefly in the closing section. AN OVERVIEW OF CHANGES IN THE BIODIVERSITY OF AUSTRALIA’S RANGELANDS There have been dramatic changes in the biodiversity of Australia’s rangelands over the last 200 years (Morton 1990; Short & Smith 1994; Smith & Quin 1996; Woinarski 2001). Patterns of biodiversity loss vary according to taxa and depending on geographic locality. Mammals of the arid and semi-arid zones have suffered most with 14 species now extinct on the mainland while the ranges of a similar number have declined markedly in extent (Morton 1990). In the 1 more mesic northern rangelands, which include the Kimberley and the tropical savannas, the situation is not as catastrophic- several declines but no extinctions in the mammalian fauna (Woinarski 2001). Rangeland birds have fared better than the mammals. Although a significant number of bird species have declined, some to the point of regional extinction, none has totally disappeared (Morton 1990; Reid & Fleming 1992; Woinarski 2001). The same appears to be true of reptiles and amphibians (Morton 1990; Woinarski 2001). Nine plant species which formerly occurred in the rangelands are now considered to be extinct on the mainland (Leigh & Briggs 1992) while a further 22 rangeland species are considered threatened (Woinarski 2001). Decline in the biodiversity of Australia’s rangelands is ongoing (Woinarski 2001). The rufous hare-wallaby (Lagorchestes hirsutus) became extinct on the mainland as recently as 1991 (Langford & Burbidge 2001). The ranges of other arid zone mammals like the greater bilby (Macrotis lagotis) (Paltridge & Southgate 2001) and the black- footed rock wallaby (Petrogale lateralis) (Gibson 2000; Copley pers. comm.) have continued to decline since that time. Recent declines have also been noted in mammals of the northern rangelands including the northern quoll (Dasyurus hallucatus) (Braithwaite & Griffiths 1996), the pale field rat (Rattus tunneyi) and the black- footed tree-rat (Mesembriomys gouldii) (Woinarski 2000). Recher and Lim (1990) and Recher (1999) have argued that, within a few decades, the loss of bird biodiversity in the rangelands will rival that currently ascribed to mammals. The processes which have wrought change to biodiversity in the rangelands are purported to include droughts and climatic change, predation by feral animals, grazing by domestic stock and feral animals, and altered patterns of fire (Morton 1990; Woinarski 2001). Changes cannot be attributed directly to any single process and are likely due to a complex interaction between two or more of these or other factors (Woinarski 2001). In this paper we focus on feral mammals (predators and herbivores) in the rangelands. We outline the threats posed by feral mammals to biodiversity now and in the future. We also examine ways in which the level of threat posed by feral mammals can be accurately gauged so that appropriate management can be applied at the correct time and scale. THREATS TO BIODIVERSITY POSED BY FERAL MAMMALS: CURRENT STATUS, FUTURE TRENDS AND MANAGEMENT 2 Habitat degradation, competition and other impacts of introduced herbivores Rabbit (Oryctolagus cuniculus) European rabbits are widespread in the arid and semi-arid rangelands where they have had a profound effect on the vegetation (Williams et al. 1995). In many areas, perennial pasture species have been replaced by annuals largely as a result of intensive rabbit grazing (Hall et al. 1964), and until recently, the recruitment of palatable shrubs and trees was suppressed by rabbits over vast expanses of the arid rangelands (Lange & Graham 1983; Foran et al. 1985; Cooke 1987). The Commonwealth Environment Protection and Biodiversity Protection Act (EPBC) (1999) identifies key processes which threaten the survival, abundance and/or evolutionary development of native species. Accordingly, Threat Abatement Plans must be prepared and implemented. Competition and land degradation by European rabbits is currently listed as a key threatening process under this act. The impacts which rabbits have, or might have had, on native mammals is open to conjecture. Several authors (Morton 1990; Williams et al. 1995; Woinarski 2001) suggest that rabbits have played a key role in the demise of arid zone mammals whether directly through competition and habitat degradation or by supporting high populations of introduced predators. A recent study (Robley et al. in press) indicates that the latter of these mechanisms has had by far the greater impact on native mammals. Australia wide, biological control has had a significant impact on rabbit populations. Myxomatosis, which was deliberately introduced in the early 1950s, had a marked initial impact on rabbit populations but did not solve the rabbit problem in the longterm (Coman 1999). Rabbit Haemorrhagic Disease (RHD), which became established in 1995, has had a more profound effect. It has reduced rabbit numbers across much of the arid and semi-arid rangelands by over 80% (Cooke 1999; Neave 1999). This has been enough to allow the regeneration of many perennial shrubs and trees (Sandell & Start 1999) that were threatened with extinction across the arid rangelands (Woinarski 2001). Rabbit Haemorrhagic Disease is endemic in rangeland rabbit populations and at this point in time there is no evidence of any reduction in its impact (Cooke 1999). Goat (Capra hircus) Feral goats are common in the arid and semi-arid rangelands of New South Wales, Queensland, South Australia and Western Australia- particularly where stock water has been 3 provided and dingoes (Canis lupus dingo) are controlled (Parkes et al. 1996). In 1993 the population of feral goats was estimated at 2.6 million with most of these in the rangelands (Parkes et al. 1996). Left unchecked, feral goat populations have the potential to reach much higher densities than currently exist (Parkes et al. 1996). Although the environmental impacts of feral goats are not well documented, competition and land degradation by feral goats is listed as a key threatening process under the EPBC Act. Goats are believed to have a deleterious effect on perennial vegetation, contribute to soil erosion and compete with native mammals for forage (Parkes et al. 1996; Woinarski 2001). Goats are widely harvested for commercial purposes (Parkes et al. 1996). Harvesting is usually accomplished by trapping and mustering. Alternative control techniques are ground-based and aerial shooting (Parkes et al. 1996; Edwards et al. 1997). In South Australia, yellow- footed rock-wallaby (Petrogale xanthopus) populations have responded positively to a combination of fox and goat control in recent years (de Preu et al. 2001). Horse (Equus caballus) and donkey (E. asinus) Feral horses are patchily distributed within the rangelands (Dobbie et al. 1993) whereas the major concentration of donkeys is in northern Australia (Wilson et al. 1992). The most reliable population estimates are from the Northern Territory where recent aerial surveys show that there are about 265,000 horses and 165,000 donkeys (K. Saalfeld, Parks and Wildlife Service of the Northern Territory, unpublished data, 1986-2001). Although the environmental impacts of feral horses are not well documented, it is believed that they contribute to erosion, damage vegetation and disperse weeds (Dobbie et al. 1993). While unequivocal data on the competitive impacts of horses on native animals are lacking, there has been a striking recovery in populations of the black- footed rock-wallaby on Finke Gorge National Park near Alice Springs since large numbers of feral horses were removed during the 1990s (Matthews et al. 2001). Trapping, mustering and aerial shooting are the best methods for humanely controlling feral horses and donkeys over large areas (Dobbie et al. 1993). Camel (Camelus dromedarius) Feral camels are widely distributed in the arid and semi-arid rangelands of Western Australia, South Australia and the Northern Territory. The most reliable population estimates are from the Northern Territory where recent aerial surveys show that there about 200,000 camels (K. 4 Saalfeld & G. Edwards, Parks and Wildlife Service of the Northern Territory, unpublished data, 2001). This extrapolates to a population of about 740,000 camels Australia wide (Short et al. 1988). The Northern Territory population more than doubled between 1993 and 2001 (Wurst & Saalfeld 1993; K. Saalfeld & G. Edwards, Parks and Wildlife Service of the Northern Territory, unpublished data, 2001) and, left unchecked, will double again in about 8 years. Although the environmental impacts of feral camels are not well documented, anecdotal evidence indicates that they contribute to erosion, damage vegetation and foul waterholes (P. Latz, pers. comm.). Like goats, camels are viewed as a resource and are liveharvested in central Australia. However, current offtake (approximately 5,000 p.a., P. Seidel pers. comm.) is unable to contain population growth even within the Northern Territory. Trapping and mustering are the main methods used to harvest camels for commercial purposes. In remote areas where it is not economically feasible to harvest camels, the only option available for reducing camel numbers and impacts is aerial shooting. Water buffalo (Bubalus bubalis) Feral water buffalo are confined to the wetter parts of the northern rangelands. Prior to the national Brucellosis and Tuberculosis Eradication Campaign, which saw widespread elimination of populations, there were approximately 340,000 water buffalo in northern Australia (Bayliss & Yeomans 1989a,b). The current population is about 73,000 (K. Saalfeld, Parks and Wildlife Service of the Northern Territory, unpublished data, 1997-2000). Obviously the population has enormous growth potential. Disturbance by water buffalo facilitates salt-water intrusion in wetland areas leading to wholesale vegetation change (Whitehead et al. 1990). Water buffalo have also been linked to declines in fauna populations (Braithwaite et al. 1984). Given the nature of the habitat which they occupy, aerial shooting is the most cost effective option for reducing water buffalo populations. Pig (Sus scrofa) There are estimated to be between 3.5 and 23.5 million feral pigs in Australia with most of these in the rangelands (Choquenot et al. 1996). Feral pigs occur patchily around wetland areas of the dryer rangelands but are widespread and common in the wetter northern rangelands. Pigs are omnivorous and consume a wide range of plants and animals (Choquenot et al. 1996). Pigs also root up ground contributing to soil erosion and bank destabilisation, and 5 locally threaten some plant species (Russell-Smith & Bowman 1992; Choquenot et al. 1996). Accordingly, predation, habitat degradation, competition and disease transmission by feral pigs is listed as a key threatening process under the EPBC Act. Aerial shooting, ground shooting, trapping and poisoning are among the methods used to control feral pigs (Choquenot et al. 1996). Predation Cat (Felis catus) Feral cats are distributed throughout the rangelands (Dickman 1996). Although densities as high as 6.3 km-2 have been recorded in the Mitchell grass downs during an eruption of the long- haired rat (Rattus villosissimus) (G. Edwards, Parks and Wildlife Service of the Northern Territory, unpublished data, 1994), densities in the order of 0.1-0.6 km-2 are more typical of arid rangelands (Jones & Coman 1982; Edwards et al. 2001). Contrary to the predictions of Newsome et al. (1997), feral cats have not declined in abundance in concert with rabbits over large parts of the arid rangelands in the wake of RHD (Edwards et al. in press). Predation by feral cats is listed as a key threatening process under the EPBC Act. The clearest evidence that cat predation can have a serious impact on populations of native animals comes from recent attempts to reconstruct rangeland mammal assemblages. A reintroduction programme for the rufous hare-wallaby in central Australia failed due to predation by feral cats (Gibson et al. 1994). More recently, predation by feral cats has hampered attempts to reintroduce the burrowing bettong (Bettongia lesueur) and numbat (Myrmecobius fasciatus) in Western Australia (Christensen & Burrows 1995; Friend & Thomas 1995) and the brush-tailed bettong (Bettongia penicillata) in New South Wales (D. Priddel, NSW National Parks and Wildlife Service, unpublished data, 2002). Although feral cats have been eradicated from islands using a combination of techniques (Veitch 1985; Berruti 1986; van Rensburg et al. 1987; van Rensburg & Bester 1988; Bloomer & Bester 1992), broadscale control on the Australian mainland has proved problematical (Christensen & Burrows 1995). Because feral cats rarely scavenge (Bayly 1978; Paltridge et al. 1997), it appears that successful control can only be achieved by baiting at times of low prey abundance (Short et al. 1997). Red fox (Vulpes vulpes) 6 Foxes are distributed throughout the arid and semi-arid range lands (Saunders et al. 1995). Reported densities are in the range 0.6-2 km-2 (Marlow 1992; Saunders et al. 1995) with densities being higher in areas with rabbits but without dingoes (Saunders et al. 1995). Foxes appear to be expanding their range in a northerly direction. They were rarely encountered in the Tanami Desert northwest of Alice Springs in the 1970s and early 1980s (Bolton & Latz 1978; Gibson 1986). Now they are relatively common there as far north as Tennant Creek (Paltridge & Southgate 2001). Foxes have not declined in abundance in concert with rabbits over large parts of the arid rangelands in the wake of RHD (Sandell & Start 1999; Edwards et al. in press). There is abundant evidence that predation by foxes is a major threat to many species of native fauna (Burbidge & McKenzie 1989). Studies in Western Australia have shown that foxes have a major impact on existing populations of black-footed rock-wallabies (Kinnear et al. 1988, 1998), brush-tailed bettongs (Saunders et al. 1995), numbats (Friend 1990) and tammar wallabies (Macropus eugenii: Saunders et al. 1995). Populations of all these animals increased following intensive fox control. Similarly, in New South Wales, malleefowl (Leipoa ocellata) survival has been shown to increase following fo x removal (Priddel 1989; Priddel & Wheeler 1990). In the Northern Territory, one of the two last known wild populations of the rufous hare-wallaby was exterminated by a fox (Lundie-Jenkins et al. 1993). Predation by foxes is appropriately listed as a key threatening process under the EPBC Act. Remaining wild populations of the bilby may be at risk of extinction if foxes continue to push northwards into the bilby’s remaining stronghold (Newsome et al. 1997). Foxes readily scavenge and can be effectively controlled over large areas using baits containing compound 1080 (sodium mono- fluroacetate) (Christensen & Burrows 1995; Saunders et al. 1995; Thomson et al. 2000). MONITORING THE IMPACTS OF FERAL MAMMALS Designing a monitoring program Monitoring and management action Population monitoring serves two purposes in vertebrate pest management. Firstly, it provides necessary information, such as population size or trend, which underpins management-based decision making (Elzinga et al. 2001). Secondly, it provides ‘research and performance evaluation’ (Possingham 2001), by indicating whether a control strategy is working and 7 allowing that strategy to be refined. This latter role forms the basis of adaptive management, where management learns about the system through monitoring responses to management actions (Shea et al. 1998). Ideally, adaptive management should be structured like an experiment, with controls and a range of treatments, in order to maximise what can be learnt from manipulating the system. Ideally, it is pest damage that should be monitored (Hone 1994). However, if it is impractical to monitor impacts or if the relationship between damage and absolute population size (N) of the pest is known, then one can monitor N and use it to indicate damage. Possible relationships between population size and damage are shown in Fig. 1. Figure 1 is an oversimplification, particularly as it assumes that damage (e.g. overgrazing) will be a function of pest density only, and ignores temporal and spatial variation in environmental conditions and variation in the density of other species. For example, in the rangelands, the potential for land degradation through overgrazing is likely to be heightened during drought. Drought would shift the impact curve from C to A in Fig. 1. The cost of controlling large herbivores is also likely to be cheaper during drought as animals are concentrated around persistent feed and water. Whether or not there is a linear relationship between pest abundance and impact (B in Fig. 1) will vary case-by-case (Hone 1994). Nonlinearities may signify the existence of stable states that can provide a level of resilience to a change. This may be vegetation resisting further degradation or rehabilitation (Friedel 1991), or a population of endangered prey remaining trapped within a ‘predator pit’ despite some effort expended in control (Pech et al. 1995). 8 Cost ($) Impact (A) Impact (B) Control Impact (C) Pest density Figure 1. Relationships between pest density and the per capita cost of control (dashed line) and the per capita cost of damage or impact (solid lines). A target density could be identified at the point of intersection. After Braysher (1993). Dealing with uncertainty Uncertainty (or error) in our understanding of how a rangeland system functions, and how it will respond to management intervention (e.g. predator control), arises from a number of sources. Process uncertainty results from natural variation in the environment through time and can be further broken down into demographic and environmental stochasticity (Hilborn & Mangel 1997). Observation uncertainty results from measurement or sampling error (Hilborn & Mangel 1997). Finally, where mathematical models are used, there is model uncertainty, which arises from ignorance of how a system works, and includes uncertainty in parameters and in the actual model structure (Parma et al. 1998). There are a range of ways in which observation uncertainty can be reduced including increasing sample size and stratification. The appropriate level of precision should be identified before a survey is undertaken and will depend on factors that include the size of the population change that needs to be detected (sensitivity sensu Eberhardt 1978) and/or the proximity to some threshold that will trigger a management action. 9 Process uncertainty introduces complications when interpreting a time series of data, as important trends may be obscured by inherent variability in the ecological processes involved. Detecting trends in the face of process uncertainty requires an understanding of how populations respond to environmental change through time, primarily food supply driven by rainfall. Estimates of precision from a regression of ln(N) against time will be misleading because data are likely to be autocorrelated (McCallum 2000). An important problem is determining how much monitoring should be done. Should more or fewer resources be invested in estimating vertebrate pest numbers? In wildlife management, little use is made of standard errors in broad-scale population estimates beyond assessing whether there has been a significant year-to-year change in population size. This uncertainty in population size (observation error) can also be used to determine the risk of an undesirable management outcome. This may be excessive control, leading to wasted effort, or too little control, leading to less effective damage mitigation. The uncertainty in future population size due to environmental stochasticity (process error) and model uncertainty would also need to be incorporated into this risk assessment. This approach is well established in conservation biology (e.g. Burgman et al. 1993), but can be applied to other areas of wildlife management (e.g. McCarthy 1996). This framework could then be used to determine the adequacy of present monitoring. Surveys could obviously be increased in frequency or intensity to reduce risk, or reduced in frequency if the resulting increased risks were acceptable. Indices of abundance and bias An index I of population size (or density) is some attribute that changes in a predictable manner with changes in absolute population size N (or density) (Caughley & Sinclair 1994). I may be a count of sign (e.g. the density of tracks), which indicates that the species of interest is present, or a direct count of individuals of the species (e.g. the number of kangaroos seen per km along a road). Strip-based direct counts of animals are best viewed as indices of population size as they typically underestimate N because observers do not see all of the individuals (ie. the counts are negatively biased) (Jolly 1969). Indices are useful for tracking relative abundance only if the ‘proportionality constant’ β connecting them to N (i.e. I=βN) remains more or less constant among surveys (Lancia et al. 1996, McCallum 2000). This is often assumed but rarely tested. Factors affecting counts (e.g. weather, observer experience, velocity of travel) will therefore need to be either standardised or corrected for to ensure β 10 does not vary. Procedures for determining bias in direct counts are described in the next section. For incomplete counts, observation error comprises sampling variation and error in β. Using the delta method (Seber 1982), an approximation of the variance of N is (Lancia et al. 1996): var( x ) var( β ) var( N ) = N 2 2 (1 − α ) + β2 x where x is the raw count of animals, α is the proportion of the study area sampled (i.e. 1-α is the finite population correction), and β is the proportion of animals counted in the sample unit, such as an aerial survey strip (i.e. N = x ). αβ Current ‘best practices’ Across the rangela nds, little attempt is made to monitor the actual impacts of feral mammals. There may be several reasons for this. First, the impacts of many of the feral mammals that inhabit the rangelands are poorly understood (see above). It is unfortunate that many of the studies which have addressed this issue lack scientific rigour (Hone 1994). Second, most feral mammal species are widely distributed and it is neither practical nor efficient to monitor impacts at the appropriate scales. Third, grazing impacts in particular are often confounded due to the presence of other grazing species. As a result, monitoring, where it occurs at all, usually involves population assessments. These are used as the basis for management decisions, the untested assumption being that higher densities equate to higher impacts. Population estimates for larger herbivores tend to be undertaken at irregular intervals and it is important to consider the intervening environmental conditions (ie. process error), in addition to observation error, before invoking long-term trend. For large feral herbivores (horses, donkeys, camels, goats, buffalo), broad-scale monitoring can be achieved efficiently through aerial survey, using either fixed wing aircraft or helicopters and either strip or line transects (Bayliss & Yeomans 1989a,b; Wurst & Saalfeld 1994; Southwell 1996; Pople et al. 1998). Aerial survey data incorporate two types of biasavailability bias and perception bias. Perception bias is a result of observers missing animals that are potentially visible, while availability bias arises because some animals are concealed from the observers (Marsh & Sinclair 1989). Survey parameters (e.g. height, velocity, strip 11 width) have varied among surveys although comparisons can usually be made over time and between areas when β, and therefore N, has been estimated. In aerial strip transect sampling, factors affecting bias (e.g. observer, group size and habitat) can be identified using double counting and log- linear modelling (Marsh & Sinclair 1989; Choquenot 1995b; Pople et al. 1998). An assumption of double counting that is potentially violated in aerial surveys is equal ‘catchability’ of sightings. Caughley & Grice (1982) suggested through simulation that violation of this assumption only introduces significant bias when sighting probabilities are <0.5. Unequal catchability, or capture heterogeneity, is often a problem in mark-recapture studies, and can be partly addressed in aerial surveys by modelling the heterogeneity. For example, the relationship between sighting probability and group size can be quantified using regression methods (Graham & Bell 1989; Pople et al. 1998). The potential overall bias can also be tested with independent data, such as index- manipulationindex (Bayliss & Yeomans 1989a). Aerial line transect sampling offers survey-specific correction for perception bias (Buckland et al. 1993). The method has generally been limited to helicopter surveys, because the primary assumption of sighting probability on and near the line being 100% is more likely to be met than in fixed-wing surveys. However, combining line transect sampling and double counting (Borchers et al. 1998) can account for uncertain sighting probability on the line. This may allow wider application of the technique to fixed-wing surveys, which are considerably cheaper than surveys by helicopters. Feral pigs are cryptic, and although changes in absolute density may be monitored by markrecapture (e.g. Caley 1993), over large areas pigs are most effectively monitored by aerial survey. Due to the dense cover typically occupied, surveys are conducted at low altitude (30.5–46 m) and low velocity (80–93 km h-1 ), necessitating the use of helicopters (Hone 1990a,b; Choquenot 1995a). Surveys are generally conducted in the cool of early morning and evening, when a greater proportion of pigs occur in more open habitats. Spotlight transect counts are the most popular method of monitoring rabbit populations, the data serving as indices of relative abundance. Line transect sampling (Buckland et al. 1993) could potentially give estimates of absolute abundance, providing the proportion of rabbits underground (=availability bias) is estimated. In practice, spotlight counts appear to reliably 12 detect changes in rabbit abundance arising from active management such as poisoning (Robinson & Wheeler 1983), predator removal (Newsome et al. 1989), or introduction of disease (Mutze et al. 1998). Although there have been concerns expressed as to the precision and accuracy of spotlight counts (Fletcher et al. 1999), Caley & Morley (2002) demonstrated that properly conducted spotlight transect counts are highly precise. For cats and foxes, broad-scale monitoring is likely to be limited to presence-absence or abundance rank. Track-based methods appear most appropriate as they offer reasonable precision and are time efficient (Edwards et al. 2000). There are two classes of track-based techniques- active and passive. Active methods use attractants (e.g. Linhart & Knowlton 1975; Roughton & Sweeney 1982; Conner et al. 1983; Smith et al. 1994; Thompson & Fleming 1994); passive methods do not (e.g. Newsome et al. 1972; Thompson et al. 1989; Allen et al. 1996; Mahon et al. 1998). Such indices have a non- linear relationship with density, with the index becoming less sensitive as density increases (Edwards et al. 2000). This may be acceptable if changes in the lower density range need to be detected, as is often the case in the arid rangelands (Edwards et al. 2000). New approaches Thermal imaging has been used in the survey of a number of large vertebrate species since the late 1960’s, primarily in temperate habitats (Croon et al. 1968; Graves et al. 1972; Parker & Driscoll 1972; Wride & Barker 1977; Wyatt et al. 1980, 1985; Best et al. 1982; Trivedi et al. 1982; Wiggers & Beckerman 1993; Boonstra et al. 1994; Garner et al. 1995; Naugle et al. 1996; Gill et al. 1997; Havens & Sharp 1998; Focardi et al. 2001). Early surveys had limited success (Wiggers & Beckerman 1993), due primarily to poor resolution and temperature differentiation of the imaging equipment. Interference from canopy cover, in particular, led to difficulty in distinguishing between animal and background (Croon et al. 1968; Graves et al. 1972; Parker & Driscoll 1972). Additionally, the cost of thermal imaging equipment was a significant impediment to the broad-scale application of the technique (Croon et al. 1968; Graves et al. 1972; Parker & Driscoll 1972). Recent advances have resulted in substantial improvements in the accuracy of thermal imagery for wildlife survey. It is now generally accepted that the technique detects a greater proportion of the population under investigation than other methods (Gill et al. 1997; Havens 13 & Sharp 1998; Focardi et al. 2001). For example, on Cobourg Peninsula, Northern Territory, approximately four times as many Bali cattle (Bos javanicus) were detected with thermal imagery than with standard aerial survey (K. Saalfeld & D. Lawson, Parks and Wildlife Service of the Northern Territory, unpublished data, 2001). However, thermal imaging remains expensive. Depending upon the intended use (e.g. ground verses aerial survey) and the desired level of resolution, thermal imaging equipment can cost from $100,000 to $500,000 (FLIR Systems Inc, Sierra Pacific InfraRed). In addition, expensive survey platforms (e.g. helicopter) are required as surveys need to be conducted at relatively low velocities. 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