Theme edition. The dilemma: cull, contain or conserve, edited by Thomas Newsome, Chris Dickman and Daniel Lunney. The important role of private wildlife organisations in facilitating scientific research: A case study of the Australian Dingo Foundation Bradley P. Smith1,2, Shennai G. Palermo2, and Lyn Watson2* 1 Smith Human-Wildlife Coexistence Lab, School of Human Health and Social Sciences, Central Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 Queensland University , Adelaide Campus, 44 Greenhill Road, Wayville, , Australia. 2 Australian Dingo Foundation, Toolern Vale, Victoria, Australia. * To whom correspondence should be addressed: Mrs Lyn Watson. Australian Dingo Foundation. P.O. Box 502, Gisborne, Victoria 3437, Australia. Tel: +61 03 5428 1245. E-mail: [email protected]

As we enter an era of global mass extinctions, it is important to tackle wildlife research and conservation from multiple fronts, including those made available by wildlife organisations, zoos and sanctuaries. Captive studies are particularly useful when studying free-ranging populations is difficult, and/or when controlled conditions are required. Yet, despite the significant role that they play in supporting research and conservation of species and ecosystems, they are rarely recognised in the scientific literature. Here we present a case study of the Australian Dingo Foundation (ADF), a private organisation and captive breeding facility that actively supports research and conservation efforts relating to the dingo ( dingo). Over the past decade (2010 to 2020), the ADF has facilitated research across eight research disciplines that include archaeology, behaviour, biology, cognition, evolutionary psychology, non-lethal management, reproduction and parental behaviour, and vocalisations. This has resulted in at least 21 published scientific studies which are summarised in this paper. As this case study demonstrates, captive facilities have the potential to contribute to the understanding and conservation of dingoes by providing practical alternatives to, and/or supplement studies of free-ranging populations. We conclude by outlining some of the implications and limitations of conducting research using captive dingo populations. ABSTRACT

Key words: Science; research; conservation; captivity; zoos; sanctuaries; organisations; not-for-profit.

DOI: https://doi.org/10.7882/AZ.2020.023

Introduction Not-for-profit wildlife organisations, zoos and sanctuaries behaviour. Work was directly carried out at their custom provide a fundamental role in the conservation of research facilities in Alice Springs, NT (Division of Wildlife many of Australia’s native species such as the dingo Research; Figure 1) and Canberra, ACT (Arid Zone (Canis dingo). Not only do captive populations provide Research Institute). Funding for much of the early research opportunities to observe and engage with wildlife, with dingoes, including those carried out by the CSIRO, they are actively involved in conservation education, was provided by the Australian Meat Research Committee wildlife tourism, breeding programs, and the facilitation and the Australian Meat and Livestock Corporation, in an and financial support of research (Tribe 2001; Tribe effort to “know thine enemy” (Newsome et al. 1972, p.1), and Booth 2003). The contribution to the latter is and thus assist with eradication attempts. Contemporary particularly important, because research investigating funding for dingo research comes from a range of sources, the biology and behaviour of many free-ranging species and is conducted by scientists from around the world. These is rarely possible, or at least very difficult. researchers represent a number of scientific disciplines, and assert that the dingo is a scientifically and culturally There is a long history of utilising captive dingoes for valuable species (Smith 2015). Research and the number research, the first of which were carried out by CSIRO of scientific publications focusing on the dingo has also in the 1970s, 80s and early 90’s (Corbett 2001). These increased dramatically over the past few decades (see Smith captive studies, led by ecologists Alan Newsome, Laurie and Appleby 2015), including research involving captive Corbett, and Peter Catling, were used to complement dingo populations. their field studies on several aspects of dingo biology and Australian 2020 Zoologist A Smith et al. Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021

Figure 1. CSIRO experimental officer with captive dingoes held at their research center in the Northern Territory. Photo taken in 1967. Source: National Archives of Australia, A1200, L64195. The benefits of captive research In captivity, not only can housing conditions be manipulated and replicated, but the identification and Like most wild canids, dingoes are notoriously difficult to selection of individuals is easier (including sex, age, body study in the wild because they are highly mobile, occupy mass, ancestry, genetics). Scientists have greater power large territories, are difficult to see in many environments, to take medical, physiological or observational samples and are generally reclusive (Klinghammer and Goodman from individuals with known life histories. To give an 1987; Morrant 2015). As such, the study of many aspects example, to estimate the food consumption of dingoes, of biology and behaviour in the field become impractical Green (1978) required carefully controlled experiments and are better suited to captive situations where with tame dingoes in laboratory settings. In the CSIRO conditions can be controlled. In fact, a more complete facilities, researchers were able to extensively study dingo understanding of behaviour can often be achieved when reproductive biology, behaviour and seasonality (Newsome field observations are supplemented by, or aided by, et al. 1973; Catling 1979; Catling, Corbett, and Newsome observations conducted in captivity. To give an example, 1992); to look at the impact and identification of dingo- Allen (2015) used dingo paw prints left in sand plots to hybrids, including genetics and coat colour (Newsome determine the impact of poison baiting programs on the and Corbett 1982); and to develop age determination abundance and age distribution of the resident dingoes. techniques and measure growth rates (Catling, Corbett, To facilitate this research, measurements of dingo paws and Westcott 1991). from captive dingoes of known ages were collected and used as a reference sample. Much of our understanding Captive environments also offer a valuable and cost- of the social behaviour of wild canids was gained through effective opportunity to test the behavioural responses captive (Schenkel 1947) and later supported by of dingoes to non-lethal tools or strategies before large field studies by Mech (1970). Similarly, a lot of early scale field testing. For instance, the effectiveness of an knowledge of dingo social behaviour was based on studies ultrasonic device as a deterrent (Edgar et al. 2007). of captive populations and substituted with observations Captive dingo populations have been used to test the of free-ranging dingoes (Corbett and Newsome 1975; behavioural responses of dingoes to dingo urine (Robley Corbett 1988). Captive research continues to add value et al. 2015), and the anti-predator reactions of kangaroos to our understanding of wild canids (e.g. Kershenbaum et to dingo urine (Parsons and Blumstein 2010). The al. 2016; Gese et al. 2018). collection of urine (including from known individuals) for such purposes would be virtually impossible in the Australian B Zoologist 2020 The contribution of captive dingo populations to scientific research wild from living , as would a controlled setting for as government watchdogs and lobbyists. 5) Facilitate determining its effect on behaviour. scientific research. The significance of these roles is largely under-acknowledged, particularly by the scientific There are many questions regarding ethology and community. With this in mind, we wish to draw attention psychology that require well-socialised animals that can to some of the positive and valuable roles that sanctuaries only be found in captive situations. Some argue that can play in aiding scientific studies and conservation efforts. for certain behaviours (such as following social cues), One organisation, the Australian Dingo Foundation (and wolves must be intensely socialised and kept separate its affiliated Dingo Discovery Sanctuary and Research from conspecifics for most of their first 4-6 months in Centre), has provided an invaluable contribution to the

order to get accurate results (suggested by Klinghammer scientific understanding of the dingo across multiple fields Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 and Goodman 1987, and adopted in the study by Miklósi of research. et al. 2003). Research investigating dingo cognition and behaviour conducted by Smith and Litchfield (2010a; The Australian Dingo Foundation (ADF), established 2010b; 2013) and parental behaviour (Hudson et al. in 2006, is a not-for-profit organisation and Federally 2016; Hudson et al. 2019) were possible because captive registered conservation charity. The ADF supports a populations provided a sample of individuals who were captive population of dingoes at the Dingo Discovery known, who were comfortable being in test situations Sanctuary and Research Centre (hereafter Dingo and handled by staff, and a controlled environment for Discovery Centre). The Dingo Discovery Centre, the experimental setup. In addition, captivity presents established in 1990, is located on 40 acres in the foothills opportunities for ad hoc observations to occur, such as the of the Macedon Ranges, Victoria (Figure 2). At any one use of tools (Smith et al. 2012). time, the sanctuary houses around 40 genetically ‘pure’ and entire adult dingoes in custom-built enclosures. This Socialised dingoes also aid the process of taking biological includes a population of captive and wild born individuals samples in a stress-free manner, often avoiding the need for from all regions of Australia. For a comprehensive summary sedation. An example of this was the longitudinal study by of the housing and management of this population see Smith, Flavel, and Simpson (2016) who measured seasonal Smith and Watson (2015). variation in cortisol. Not only does interacting with captive animals provide insights not otherwise practical The ADF and its volunteers play a significant role in in the wild (e.g., comparative studies of problem solving dingo conservation through advocacy (e.g., organising behaviour - Smith and Litchfield 2010a, 2010b, 2013), petitions, lobbying government, social media presence), but failure to socialise captive wolves and dingoes for education (e.g., running sanctuary tours, puppy open days, certain experiments may lead to erroneous results because fundraising events, community and school education the presence of humans, particularly in close proximity, sessions), and an active breeding program (e.g., maintaining might disrupt the ’s behaviour (Klinghammer and several breeding lines, supplying zoos and wildlife parks Goodman 1987). with dingoes for exhibition). Through these activities they also facilitate and support non-invasive scientific Through wild rescue and captive breeding programs, research by making their captive dingo population populations held in private sanctuaries may provide a available to national and international scientists. The reservation of genetically ‘pure’ dingoes from regions Dingo Discovery Centre has been directly supporting around the country, should dingoes become endangered genetic studies since its inception, and behavioural studies or absent. They may also be useful for reintroduction since the late 1990’s (Wilton et al. 1999; Wilton 2001), programs involving the release of dingoes into national and the ADF has supported dingo research by providing parks (Ritchie et al. 2012; Newsome et al. 2015). Private financial support through scholarships such as the ‘NWG sanctuaries with active breeding colonies, typically have and Anne Macintosh Student Scholarship’. skilled dingo handlers, who can likely offer valuable services and consultation for reintroduction and rewilding programs. The sanctuary maintains a healthy, entire, and diverse population of dingoes which enables them to suit the requirements of researchers from a plethora of research The Australian Dingo Foundation disciplines (for examples, see Figure 3). These include (but There are a handful of private not-for-profit organisations not limited to): animal behaviour (e.g., social behaviour scattered across the country that maintain captive dingoes and parental care), cognition (e.g., problem solving, and are dedicated to dingo conservation. As outlined higher order behaviour), evolutionary and comparative by Smith and Watson (2015) these organisations make psychology, biology (e.g., microbiome, stress responses, at least five key contributions. 1) The preservation of urine chemistry), non-lethal management (e.g., testing genetically ‘pure’ dingoes from regions across Australia efficacy of non-lethal biological deterrents), and genetic (i.e., dingoes with no modern dog ancestry). 2) Provide testing. To demonstrate this, in Table 1 we present a source of animals for displays at wildlife parks, and for a number of published scientific studies that resulted future rewilding programs. 3) Raise community knowledge from research that took place or based on dingoes held and awareness of dingoes and their conservation. 4) Act at the sanctuary between 2010 and 2020. The list of Australian 2020 Zoologist C Smith et al.

A) B) Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021

C)

Figure 2. The Dingo Discovery Centre. (a) A section of the custom dingo housing (Photo: Bradley Smith). (b) Closeup of the enclosures (Photo: Bradley Smith). (c) A dingo in one of the exercise yards (Photo: Shari Trimble).

publications was sourced from the records maintained by physiological studies of dingoes that are born and raised the ADF, and the methodology and acknowledgements of in captivity (O’Regan and Kitchener 2005; Hartstone- each publication cross-checked to ensure the sanctuary Rose et al. 2014), particularly where selective breeding or foundation was acknowledged as a contributor. It is and multiple generations of captive-bred animals are used. likely that additional studies have been conducted at Although in saying this, the appropriateness is entirely the sanctuary, but the results not published, and it is also dependent on the purpose of the study. For example, possible that not all published outcomes are presented in captive-bred animals can provide a unique opportunity the table (although none were deliberately omitted). to assess the relative importance of environmental and genetic influences on morphology (O’Regan and Kitchener 2005); for examining human-animal relationships and Cautions when using captive dingo interactions (e.g. Ward and Melfi 2015); and of course populations for scientific studies for studying captive welfare, behaviour and management It is important when studying the behaviour of wild (Hill and Broom 2009). animals in captivity that they be kept in a way that maximises their opportunities for the expression of normal There are also some behaviours observable in captivity behaviour. One concern that has been raised about using that may not translate to wild populations. For example, in captive populations for research, and their value to dingo wild canids, tool use has been observed in captivity (Smith conservation more broadly, is that captivity somehow and Litchfield 2012) but not in free-ranging populations. changes the individual so it no longer truly represents Although this provides insight into the cognitive potential the species. Corbett (2001) for instance, argued that a of the species, the behaviour observed in captivity is likely captive dingo ceases to be a dingo and is “just another to be a reflection of increased opportunity and energy to dog” because selection pressure for certain traits can explore and manipulate objects within the environment. easily take place, both deliberately and inadvertently, More realistic and ecologically valid examples of tool-use and therefore could override the of wild and higher order behaviours have been observed in the characteristics. The impact of captivity, which include field in other canids (see Smith, Appleby, and Litchfield unnatural diets (even when foods are mimicked to provide 2012 and references within), but such observations remain adequate nutrition) and restriction of movement, is likely rare and difficult to capture. to be most significant in relation to morphological and Australian D Zoologist 2020 The contribution of captive dingo populations to scientific research

Table 1. A list of publications grouped across eight discipline areas, that has resulted from research carried out at the Dingo Discovery Sanctuary and Research Centre (DDC) between 2010 and 2020. A brief summary of the background and/or aims, methodology, and main findings from each publication are provided.

Discipline Title Background/Aims Methods Findings Reference

Archaeology Dingo scat-bone To address the lack Scats from 25 captive Wild and captive dingo skeletal Reynolds, ‘signature patterns’: of data on carnivore dingoes from the DDC modifications were similar but Dortch, An actualistic study modifications to skeletal and 31 wild dingo dependent on prey consumed. and Balme and comparison of fragments with focus on scats were analysed Captive dingo scat contained (2016) wild and captive scat- the dingo. for skeletal fragments. fewer and smaller skeletal

bone assemblages Useful as a potential Tooth ‘pits’ and ‘scores’ fragments than wild dingo Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 and interpretation of investigative tool were also measured on scats. Both tooth-mark and bone fragments from at Holocene skeletal fragments. scat techniques are best used Witchcliffe Rock Shelter, archaeological sites. in combination. Dingoes have south . characteristic tooth-marks, of which were identified on some skeletal fragments found in Witchcliffe rock shelter. Analysis of pit and Comparative approach DDC was used to Devils and tiger quolls Koungoulos, score tooth-mark sizes for archaeological pilot the methodology produced very large tooth Faulkner, from bones modified material, using dingoes, used in the larger study. marks in relation to their and by Holocene Australian devils, quolls, monitor Experimental feeding body size. Aussie fauna does Asmussen terrestrial fauna in lizards, and human trials of captive animals, not conform to international (2018) relation to body size. pit marks and scores including analysis of trends, that is, body weight left on bone. 2895 tooth marks. does not correlate with tooth mark size. Carnivores that consume bone in their diet will have specific adaptations that will determine pit mark and score sizes left on bone fragments. Behaviour The function of play To investigate the role Camera footage was The behavioural function Byosiere et bows in Canis lupus of domestication on the taken of 5 litters of play bows (i.e. as a al. (2018) and its variants: A function of behavioural (10 puppies total) visual signal) is similar for comparison of dingo play bows in canids. interacting at the DDC. dingoes, domestic dogs (Canis lupus dingo), dog Play behaviour was and wolves. Plays bows are (Canis lupus familiaris) observed and coded more exaggerated, longer, and puppies (Canis according to five criteria and involve vocalisations in lupus). and compared with domestic dogs compared with those of dogs and dingoes and wolves. wolves. Biology The effect of age, To analyse the canid Scat samples from 101 Canid microbiome differences Flavel gender, and antibiotics microbiome at different domestic dogs and can be attributed to (2012) on the canine gut life stages, and how it from 28 dingoes at the various factors, such as age. microbiome. is affected by factors DDC were analysed Bacteroidetes/ prevotella group like age and gender. To for two main bacterial and clostridia coccoides were compare the microbe classes, bacteroidetes found in Labradors and of domestic dogs and and clostridia. dingoes at different life stages. dingoes. Quantification of To better understand Saliva samples were Cubs had higher cortisol Smith, salivary cortisol from ‘stress’ and its collected from 13 levels than adults. Adults did Flavel, and captive dingoes (Canis implications on captive adult dingoes, and 8 not differ, but males during Simpson dingo) in relation to age, and wild dingoes. To dingo cubs from the breeding season had higher (2016) gender, and breeding obtain a reference DDC across several cortisol levels than females. season. range of dingo salivary time points (across a cortisol levels. day, development and seasonality). Cognition How well do dingoes To test dingoes on a A v-shaped version of Dingoes were proficient at Smith and (Canis dingo) perform spatial problem-solving the ‘detour task’ using completing the task with fewer Litchfield on the detour task. task. Previously tested four experimental errors and shorter delays than (2010a) wolves performed conditions was tested dogs. Wolves and dingoes are better than domestic on 20 dingoes at the more capable at non-social dogs - a potential result DDC. Success, latency spatial problem solving than of domestication. and errors detouring domestic dogs. fence were measured.

Australian 2020 Zoologist E Smith et al.

Discipline Title Background/Aims Methods Findings Reference

Dingoes (Canis dingo) To test the The ‘object-choice Dingoes passed 6/9 Smith and can use human social responsiveness of task’ was tested on 7 human social cues. Dingo Litchfield cues to locate hidden dingoes to human social dingoes at the DDC to performance was more (2010b) food. cues, as they have a see how they respond akin to wolves than dogs, unique evolutionary to 9 different human suggesting domestication may history that may shed social cues. Success influence the comprehension light on canid evolution. resulted in approaching of human gestures. container being referred to by human. Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 Spontaneous tool-use: To document the Two male dingoes Observations of tool-use Smith, An observation of a cognitive abilities of two at the DDC were extend to canids. Highlights Appleby, dingo (Canis dingo) captive dingoes, and the observed and recorded the dingoes potential for and using a table to access first case of tool-use in exhibiting higher order ‘intelligent’ behaviour adds Litchfield an out-of-reach food the family. behaviours. 1) Sterling insight into their cognitive (2012) reward. moved a table in the abilities and potential. enclosure to reach an otherwise unobtainable treat. 2) Teddy opened a gate latch to get to his mate when he was separated from her. Looking back at ‘looking To find out whether A ‘rope-pulling task’ was Dingoes do not rely on Smith and back’: Operationalizing dingoes look-back tested on 12 dingoes humans for assistance during Litchfield referential gaze for for human assistance at the DDC, with a tasks in the same way as dogs. (2013) dingoes in an unsolvable when faced with solvable trial, followed A variety of ‘look-backs’ was task. an unsolvable task. by an unsolvable one. observed in dingoes but was Previously tested canids Behaviour in response not likely associated with included domestic dogs to unsolvable condition seeking human assistance. This and wolves. Domestic coded. resulted in the refinement of dogs looked back to the definition of a ‘look-back’. humans for assistance, wolves did not. Evolutionary The evolution and To investigate the An ‘inequity aversion Both canid species did not McAuliffe psychology development of expression of ‘judging task’ was tested on 72 express ‘inequity aversion’; (2013) inequity aversion. fairness’ or ‘inequity dogs and 11 dingoes not sensitive when only aversion’ in dogs and at the DDC. Unfamiliar ‘confederate partner’ dingoes. To determine human handlers and received food. May not be whether this is a unfamiliar dog or a cooperative mechanism result of a cooperative dingo ‘confederate in social canids. Looking at mechanism found in partners’ were used. human handlers was more social canids, or a result Trials involved absence/ prevalent in dogs than in of domestication. presence of food. Video dingoes. recordings were coded and analysed; sensitivity to presence/absence of food, response time and looking at human handlers. Exploring the To investigate the Three experimental No evidence for overimitation Johnston, evolutionary origins origins of learning trials used on 40 dogs in dogs or dingoes. Dingoes Holden, of overimitation: A via ‘copying’ or and 13 dingoes at the may be better at finding and Santos comparison across ‘overimitating’ in DDC. 1) Transparent aspects of the puzzle that are (2016) domesticated and non- humans. To establish puzzle box, lever not irrelevant, compared to dogs. domesticated canids. whether or not needed to obtain treat. Overimitation appears to be overimitation is a 2) Non-transparent a unique form of learning in shared learning skill with puzzle box, lever not humans. dogs, which have close needed to obtain treat. social bonding with humans. 3) Transparent puzzle box, lever needed to Dingoes used as a obtain treat. comparison to dogs.

Australian F Zoologist 2020 The contribution of captive dingo populations to scientific research

Discipline Title Background/Aims Methods Findings Reference

Uncovering the origins To explore interspecific Duration of eye contact Dingoes established eye Johnston et of dog–human eye communication using made by dingoes contact more than wolves, al. (2017) contact: Dingoes human-dingo eye was recorded over a but less than dogs. Human- establish eye contact contact. Dingoes 5-minute period with canid eye contact may have more than wolves, but were tested to gain 23 dingoes at the developed at the onset of less than dogs. further insight into DDC. Familiarity was domestication. Prolonged eye canid domestication, also tested to see if contact may have evolved given their unique eye contact changed later, especially toward familiar evolutionary history. according to whether humans, as observed in

or not the dingo knew domestic dogs. Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 the handler. Non-lethal How guardian dogs To determine how Dingo vocalisation Dingo vocalisation playbacks van Bommel management protect livestock from maremma dogs recordings (howling) and scent elicited behavioural and Johnson predators: Territorial maintain territories and urine samples were responses in maremma (2015) enforcement by inclusive of livestock. taken from dingoes sheepdogs. If ‘threat’ was maremma sheepdogs. That is, whether they housed at the DDC. inside range, they left sheep do so directly when Sound and scent tests in pursuit of ‘threat’. If ‘threat’ livestock are threatened, were carried out on was outside range, they just or indirectly (through two sheep properties barked, and scent marked. sound and scent). with 4 GPS collared Thus, protection appears to maremma sheepdogs extend beyond the immediate on each. Cameras were vicinity of livestock, so they do also set up to record maintain a larger territory. maremma behavioural responses. Dingo semio-chemicals: To assess the viability Urine collected from 6 Identification of 27 chemicals Robley et al. Towards a non-lethal of the use of semio- sexually mature males were found to be unique (2015) control tool for the chemicals, extracted at a private property. to dingoes. Dominant male management of dingoes from dominant male Semio-chemicals urine will cause a behavioural and wild dogs in dingo urine, as a non- specific to dingoes were response. Utilisation of Australia. lethal tool in dingo/wild extracted from the extracted semio-chemicals dog management. above urine samples from male dingo urine appears and used to test the to be a valuable non-lethal responses of 14 dingoes tool. at the DDC. Fladry as a non-lethal To determine how Exposed 10 dingoes at Minor behavioural reaction. Smith et al. deterrent for dingoes. captive dingoes the DDC over three Fladry may offer a cost (2019) responded to fladry (a experimental trials to a effective, temporary barrier, rope line with brightly 6x6m exclusion zone but aversive effects likely to coloured flags). lined with fladry. dissipate quickly. Automated shepherds: To test whether a novel Exposed 12 dingoes Acoustic deterrent was not Smith et al. Captive dingo acoustic deterrent and at the DDC to two effective. Inflatable effigy highly (2020) responses to sound and effigy are effective non- conditions (loud gun effective, and remained so an inflatable, moving lethal deterrents. shots, and an inflatable over multiple exposures (9/36 effigy. moving effigy). Success accessed food across trials). determined if dingoes were prevented from eating from food bowls. Reproduction Variation in To compare the Literature review used Highlighted the different Lord et al. and parental reproductive traits parental and to gather parental reproductive strategies (2013) behaviour of members of the reproductive strategies and reproductive data between domestic and Canis with of domestic and wild for all canids. Captive wild canids across multiple special attention to the canids. breeding records domains. Dingoes more domestic dog (Canis collected at the DDC akin to wild than domestic familiaris). over ten years were canids. Dingoes mate during used as part of the decreasing daylight (southern comparison between hemisphere), and wolves wild and domestic dog during increasing daylight reproductive timing. (northern hemisphere).

Australian 2020 Zoologist G Smith et al.

Discipline Title Background/Aims Methods Findings Reference

Pattern of nipple use by In relation to suckling Video cameras placed No difference in suckling Hudson et puppies: A comparison behaviours in altricial inside the den of 4 behaviour was found between al. (2016) of the Australian dingo young, previously dingo litters (12 puppies domestic dog puppies or and the domestic dog. studied domestic dog total) at the DDC. dingo puppies. No indication puppies exhibited no Suckling behaviour that the lack of suckling order suckling pattern but (including nipple use, is a trait of domestication, kittens do. Aim was to time and duration of rather reflects a difference in determine whether the suckling) was observed lifestyles between canids and difference was due to and coded according to felids.

domestication. 4 measures. Previously Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021 collected data on dog litters was also used. Diurnal pattern of There is a lack of Continuous camera Evidence of bi-parental care in Hudson et pre-weaning den visits research looking at early monitoring inside dingoes, and the lack thereof al. (2019) and nursing in breeding parental care in canids. and outside of 4 in domestic dogs that may be pairs of captive dingoes This paper aimed to breeding pairs and their attributed to a shift in reliance (Canis dingo). understand the level of litters at the DDC. on humans. Mothers spent maternal and paternal Study examined the most of their time in the den care provided to pre- behaviours (such as nursing, especially at night. weaned cubs. mother/father time in Fathers slept on top of the den, nursing time) from den, seemingly to keep guard. birth to 3 weeks. Vocalisations The bark, the howl To investigate the Dingo ‘bark-howl’ The ‘howl’ and ‘bark’ Déaux et al. and the bark-howl: acoustic structure of recordings were components were both (2016a) Identity cues in dingoes’ ‘bark-howls’ and the collected from two individually distinctive - the multicomponent calls. behavioural signal ‘bark- sanctuaries, including former to a higher degree. howls’ represent in the DDC. Wild The ‘bark’ segment may dingoes. dingoes were tested in function as an initial alert WA, with recordings to obtain the receivers (i.e. played during the day/ family member) attention. night. ‘Bark’ and ‘howl’ Individuality of this vocalisation components were may inform level of reliability acoustically measured to the receiver. ‘Bark-howls’ for individuality. may function as alarm calls, signalling to receivers to be vigilant. Concatenation of ‘alert’ To further investigate Dingo ‘bark-howl’ Dingoes were able to Déaux et al. and ‘identity’ segments the function of ‘bark- recordings were used distinguish levels of familiarity (2016b) in dingoes’ alarm calls. howls’ in dingoes. from two sanctuaries using the ‘howl’ component; Aspects studied in NSW. A familiar, they were more responsive included behavioural unfamiliar and bird to unfamiliar ‘howls’ (potential responses to level of recording was tested threat) than familiar ‘howls’. familiarity, attention on 18 dingoes at the ‘Bark’ components served levels and vigilance DDC. Camera footage to initially attract receiver’s behaviours. was taken and coded attention, regardless of according to three familiarity. Dingoes were behavioural responses; observed to always be vigilant no. of looks, movement when a ‘bark-howl’ was heard, towards speaker and supporting its function as an vigilance. alarm call.

Another example is dominant female infanticide, as and lack of opportunity for individuals to disperse) reported by Corbett (1988). Corbett (1988) asserted that gave rise to this abnormal behaviour, or at least the dominant female infanticide was the primary mechanism commonality of it being observed. To date, dominant through which dingo populations were regulated. The female infanticide has not been reported in wild dingo main benefit being that mature females that no longer had populations, and in fact, there are more examples of where young (after their pups had been dispatched by a dominant it does not occur (e.g., Smith and Vague 2017). female) retained the capacity to suckle and supplementally feed the dominant female’s dependent young, increasing One final concern when using captive populations the chances of their survival. However, dominant female for behavioural experiments is the tendency to prefer infanticide has only been observed in this single captive individuals who are suited to (or tolerate) captivity population. It is more than likely that the conditions of and human interaction. For example, they do not show the study site (limited enclosure size, increased aggression, fear response to the experimenter, equipment or testing Australian H Zoologist 2020 The contribution of captive dingo populations to scientific research

A) B) Downloaded from http://meridian.allenpress.com/australian-zoologist/article-pdf/doi/10.7882/AZ.2020.023/2645716/10.7882_az.2020.023.pdf by guest on 25 September 2021

C) D)

Figure 3. A sample of research studies conducted at the Dingo Discovery Centre. (a) Testing the response of dingoes to fladry- Smith et al. 2019. (b) Cortisol sample being taken from a 3-week-old dingo cub- Smith, Flavel, and Simpson 2016. (c) A dingo completing a problem solving task, with Dr Angie Johnston from Yale University. (d) Testing whether dingoes can follow human social gestures- Smith and Litchfield 2010b (Photo credits: Bradley Smith).

scenario. This is particularly pertinent with dingoes, not appear to have much impact on the expression of where many in captivity remain timid and exhibit naturalistic parental behaviour. Where possible however, sensitivity to novelty (neophobia) and are almost always care must be taken to limit the influence of captivity to excluded from testing (Smith 2015). Excluding such experiments, limitations acknowledged, and caution used individuals can produce cleaner data (e.g., less failures), when extrapolating findings to free-ranging populations. but can also lead to false interpretations of cognitive Ideally, all captive based studies should be complemented ability (Miklósi, Topál, and Csányi 2004). Sample sizes of and verified with studies conducted in wild populations. captive studies also remain low and tend to use samples from only one sanctuary/colony (see methods column in Table 1). Sample size limitations, and the potential for Conclusion sanctuary specific behaviours (reflective of the samples By highlighting the contribution of one organisation, history and genetics, as well as the conditions of the we have shown that a wealth of knowledge can, and sanctuary) should be acknowledged. will continue to be gained about the dingo from captive based studies across a number of discipline areas. Any Despite these important considerations, we argue that of the issues related to captive based research can the impact and influence of captivity are likely to be be mitigated with sound experimental design, and/or inconsequential to several disciplines (Smith and Watson acknowledgements of limitations where applicable. There 2015). As shown in Table 1, there are many studies that are many exciting research opportunities and possibilities can be undertaken with captive populations that have with captive dingoes that can help ensure the long- few issues or concerns relating to external validity, or term future of the species. For example, facilitating the appear skewed due to captivity. Given the difficulty in testing of effective non-lethal approaches to mitigate studying free-ranging dingoes, captive populations remain dingo-livestock conflict, which remains one of the a valuable resource for scientific studies. For example, biggest challenges to dingo conservation. We conclude observing parental behaviour in dens is difficult in the by encouraging the recognition, utilisation and support of wild (especially with replicates), but can be conducted private wildlife sanctuaries and organisations in Australia. in captivity with ease, and in many ways, and does Australian 2020 Zoologist I Smith et al.

Acknowledgements The authors wish to thank all those who have been years. None of this would be possible without them. We involved with the Australian Dingo Foundation, and also thank two anonymous reviewers and Dan Lunney for Dingo Discovery Sanctuary and Research Centre over the their valuable feedback and suggestions.

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