The thylacine’s last straw: epidemic disease in a recent mammalian extinction Robert Paddle School of Psychology, Australian Catholic University, Melbourne Campus, 115 Victoria Parade, Fitzroy, Victoria, 3065. [email protected] While anecdotal accounts exist in the literature of epidemic disease as a significant factor in recent T mammalian extinctions, harder data has not previously been presented. The statistics from the deliberate killing of thylacines as a pest species support contemporary records at the turn of the C twentieth century, of an epidemic disease in thylacines and other marsupi-carnivores. For the first A time, detailed symptoms and statistics of the disease are presented, as recorded by museum staff, and zoological-garden curators and veterinarians. It is argued that the effects of the disease in captivity, R which more than halved thylacine longevity, and preferentially affected juveniles, are conformable with T the expression of the disease recorded amongst wild thylacines, and demand a recognition of the S importance of this disease as a major factor in the thylacine’s recent extinction, and its consideration B as an influential factor on the distribution and population dynamics of extant marsupi-carnivores. It also practically demonstrates the obvious potential for disease to have been involved in megafaunal A extinctions in the past. Key words: Thylacine, Tasmanian devil, marsupi-carnivores, extinction, disease, megafauna. Introduction The suggestion that a hyperdisease, introduced by humans by anecdotal accounts of disease - comment re distressed or their commensals invading a geographic entity for and diseased individuals accompanies the extinction early the first time, may have been a significant factor in late- last century of two endemic murids on Christmas Island Pleistocene mammalian megafaunal extinctions (MacPhee (Andrews, 1909; Pickering & Norris, 1996) - harder data & Marx, 1997), warrants a careful re-examination of the has not previously been offered for a disease-associated, evidence for the involvement of disease in any recent recent mammalian extinction. mammalian extinction. Introduced disease organisms The thylacine (marsupial wolf, or Tasmanian tiger, appear to have been significant factors in the extinction Thylacinus cynocephalus) lived in stable monogamous of some gastropod (Cunningham & Daszak, 1998) and pairs, presiding over a wide-spread hunting territory. Slow amphibian (Pounds et al. 2006) species, and in some breeding and long-lived (nineteenth-century records recent avian extinctions in Hawaii and New Zealand (Groombridge, 1992; van Riper et al. 1986, Warner, 1968). suggest over 14 years in captivity1), they produced up This avian data has led discussion, debate and modelling to four young every second year. Adolescent thylacines, of the possibility of hyperdisease amongst late-Pleistocene having spent 12 to 18 months co-operatively pack hunting mammalian faunal assemblages (Lyons et al. 2004). While with their parents, finally left the family group before the recent mammalian loss has occasionally been accompanied next generation of young became semi-independent of 1 In April 1856, unannounced and unexpected, a “Mr. Martin” arrived in London at Regent’s Park Zoo with a male thylacine. It shares the record for the longest-lived captive thylacine; as London specimen iii, later on-sold to become Berlin specimen i, it survived for 103 months in European captivity (and this is the longevity figure given for the specimen in Table 5). Its April arrival, however, necessarily followed a two to three month ocean voyage, indicating that it left Tasmania in January or February 1856. The fact that it survived the journey also suggests that it was well-adapted to captivity before departure. While that is all that is definitely known of the origins of this specimen, there is a conformable history of a captive thylacine to pair with it. Ronald Campbell Gunn, living near Launceston, took a thylacine into captivity in June 1851 (which must, at a minimum, given the specimen’s survival, have been the product of a late winter breeding the previous year, and thus born around September 1850). In October 1853 he placed it under the care of Chester Wilmot who initially took the thylacine to Hobart, intending for it to accompany him on his return voyage to England. Henry Propsting, however, who had opened the first zoological garden (and museum) in Hobart, became aware of the thylacine’s presence, and wrote to Gunn, pulling out the nationalistic stops and requesting the opportunity to display the thylacine locally, rather than having it sent abroad. To this Gunn agreed, and the thylacine entered the display at Propsting’s zoo in January 1854, with the extracted promise, however, that Propsting would forward the specimen to London if or when he had no further use for it. Propsting closed his zoo, left Hobart and retired to his country estate in January 1856. Having decided in 1855 to close the zoo, he offered his museum collection (largely based on departed specimens from the zoo) to the Royal Society of Van Diemen’s Land. Many of Propsting’s museum specimens were greatly received by the Society, but there was no thylacine material in the Propsting museum collection, suggesting that the animal was still alive. In accordance with the agreement he made with Gunn, Propsting would have placed the thylacine on board ship, bound for the Zoological Society of London, around January 1856. While no archival records have as yet been found that directly link this January 1856 Hobart departure and April 1856 London arrival, circumstantial evidence points to these being one and the same specimen, and suggests that, on its death in Berlin, this thylacine was at least 14 years and two months of age. As a further point, it also needs to be recognized that nineteenth-century, captive-marsupial longevity did not reach the levels obtained later in zoological gardens in the twentieth and twenty-first centuries, as experience, husbandry and veterinary knowledge of marsupials has improved over time (Holz, 2008). Australian 2012 Zoologist volume 36 (1) 75 Paddle the pouch. Adolescent thylacines potentially travelled last three years of this eight-year time period, from June widely to establish a pair bond and their own hunting 1898 to June 1901, starting with three specimens [xxii to territory (not necessarily in that order); such behaviour xxiv], not from the north-east, but from the Western Tiers, allowing for a frequency dependant transmission of any obtained in June and December 1898 and July 1889.) epidemic disease, with even greater adolescent dispersal McGowan wrote to Melbourne Zoo in November 1906 required after the appearance of the disease within the that “Tasmanian Wolves were almost extinct & Tasmanian species. After the invasion of Tasmania by European Devils [Sarcophilus harrisii] very difficult to obtain” (letter colonists at the commencement of the nineteenth November 1906). In writing to the National Museum century, thylacines began to suffer prolonged predation; of Victoria, McGowan noted that “complete skeletons” labelled as a pest and killed both by farmers (over the from thylacines possessing “damaged or rotten skins” were rural myth of the thylacine as a significant sheep-killer available, but that fine, entire specimens of the species [Paddle, 2000]), and by snarers for the fur trade (over were now almost impossible to find (letter 21/6/1909). the annoying, but understandable, tendency for any Scott commented upon the recent difficulty of obtaining marsupi-carnivore to view a snared possum, wallaby or thylacine specimens for the museum from trappers, kangaroo as a free-lunch). offering 7/6 for the body of a thylacine, more than Casually collected anecdotal records and early bounty doubling the price to 17/6 if the skin was in a fit state analyses have, at times, prompted the suggestion that for preservation (letter 10/5/1899) – such adult skins, no disease was a major factor in the extinction of the matter the quality, having already raised for the trapper species, that occurred when the last known specimen at least £1 from the government bounty. He apologized (xxix) died in Hobart Zoo during the night of 7th to his professional colleagues for the necessity of now September, 1936.2 Representative examples of authors sending less-than-perfect specimens to other museums as claiming a disease-assisted extinction include Hickman representatives of the species - “I think it worth sending a (1955), Guiler (1961), Sharland (1971a), MacPhee damaged animal … [as] good specimens are rare and not and Marx (1997) and Paddle (2000). However, the easy to obtain” - and noting that the most recent thylacine scattered and largely anecdotal evidence previously dying in Launceston Zoo was “in poor condition and fit available allows of alternative explanation, and, on for nothing” (letter 3/8/1901). Externally, such “poor not unreasonable grounds, these claims have been condition” specimens exhibited areas of significant hair previously dismissed as wanting (Johnson, 2006). loss, skin lesions and mange. The National Museum of This paper provides the first detailed description by Victoria, anxious for any additional thylacine specimens, professional scientists of the epidemic disease in the was prepared to take such “poor condition” specimens thylacine. Data is presented on captive longevity that off the hands of the Queen Victoria Museum, and were directly relates to the bounty records and strongly pleased to note how “by exercising care, we have been suggests that the disease was a significant factor leading able to save the skin, which showed traces of mange” to the extinction of the species. (Kershaw, letter 4/6/1903).3 Scott was also aware that the disease was not just restricted Initial Disease Recognition to thylacines, but that it had also dramatically reduced The first professional scientists to recognise that something Tasmanian devil numbers (letter 25/4/1904), to the point dramatic had occurred to the thylacine in north-eastern where he was also forced to send damaged, less-than-perfect Tasmania at the end of the nineteenth century were Wm. devil specimens to other museums (letter 20/6/1904). McGowan, Superintendent of the Launceston City Park McGowan similarly recognised the problem that the skins Zoo, and Herbert Scott, Director of the Queen Victoria of diseased devils that died in the zoo presented, in the Museum, Launceston. setting-up of full mounts, “because sometimes pieces of fur get knocked off” (letter 18/6/1907). As recalled by the In the eight years from the first display of the species in farmer Lewis Stevenson (interview 1/12/1972): “devils … June 1885, to July 1893, Launceston Zoo obtained 21 got the mange like the tigers … their hair fell out and left thylacines, largely from north-eastern Tasmania. (Two the black skins bare in the bad ones”. juveniles, one obtained from Montagu in the north-west of the state in November 1885 [iii], the other from Bream Few professional scientists at the time published comment Creek in the south-east in June 1886 [v] being the only on the existence of an epidemic disease affecting marsupi- known exceptions.) However, in the next eight year carnivores, but recognition of the existence of the disease period, up to June 1901, McGowan managed to obtain is to be found in the letters passing between them, in their only six thylacine specimens. (For the first five years, daily institutional records, and in their private diaries; between July 1893 and June 1898, McGowan was unable such being the primary sources on which this paper is to obtain any thylacines, from any locality whatsoever, for based. Note, however, that Albert Le Souëf jnr, with the zoo. The six specimens referred to all arrived in the a background knowledge of marsupi-carnivore arrivals 2 Throughout this paper, individual zoological garden thylacines are identified by roman numeral alone, in both text and tables 5 and 6, indicative of the chronological order of their display. Individual Tasmanian devils mentioned in the text are identified by roman numerals preceded by “S”. These designations represent a work-in-progress, with some recently discovered specimens, falling between two previously identified individuals, temporarily identified with the roman numeral of the earlier specimen, followed by the letter “a”. 3 Unfortunately, none of these mangy skins currently remain in the Museum’s collection, as later curators and collection managers had them destroyed, on the assumption that these damaged skins were exhibiting contemporary Dermestid and/or moth attack. Australian 76 Zoologist volume 36 (1) 2012 The thylacine’s last straw at Melbourne Zoo, and Moore Park and Taronga Zoos, particular thylacine interests on Woolnorth. There was Sydney; Harry Burrell, the widely travelled naturalist and only ever one “tiger man” employed at any one time at mammalogist; Frederic Wood Jones, Professor of Anatomy Woolnorth (and this was not, in any sense of the words, at the University of Adelaide; and Ellis Troughton, a full-time position [Paddle, 2000]), and no change was Curator of Mammals at the Australian Museum, Sydney, made to the level of bounty payment offered by the did publish comment as to the existence of an epidemic Van Diemen’s Land Company; which simply matched disease on the mainland amongst the Dasyurid marsupi- that of the government bounty scheme, from which carnivores at the turn of the twentieth century (A.S. Le recompense was later recovered for the bounties paid Souëf & Burrell, 1926, p324; Troughton, 1941, p43; and by the Company. The behavioural change suggested Wood Jones, 1923, p92). would thus appear to be thylacine-based, with an increased ease of capture and killing, or the discovery Two commonly reported labels for the disease were of dead specimens, being a product of the disease. The recalled by farmers, trappers and old-timers. Explaining destruction of these 55 thylacines represented the some of the immediate internal effects of fever effective end of the species as a self-sustaining unit on (lassitude, coughing, diarrhoea and thinness), and the 150,000 acre (60,700 hectare) Woolnorth property vulgarly presented as the likely origin of the disease, is (within which the majority holding of the land lay to its description as “distemper”. (For anecdotal reference the north of Mt. Cameron). thereto see Cotton [interview 1980], Guiler [1958, From 1904 onwards, occasional adolescent immigrants 1985], Hickman [1955], Sharland [1956, 1971a, letter from the surrounding wilderness areas were snared as they 23/6/1972] and Skemp [1958].) Describing the longer- passed through Woolnorth, or attempted to settle and term, external effects of the infection (lesioned skin breed, but they were few and far between. During the next with significant hair loss and external bleeding), is ten years, 1904 - 1913, only seven thylacines were killed its description as “mange”. (For anecdotal reference or captured (one in September 1906, and an entire settled to mange see Brown [1972, 1973], Guiler [1986], family of two adults and four young in late May and early Sharland [1956, 1960, 1971a, 1971b, letter 23/6/1972] June 1909). For the ten years from 1914 – 1923 there were and Stevenson [interview 1/12/1972].) no thylacines killed or captured at Woolnorth. In the next Alongside these contemporary written accounts by decade, 1924 – 1933, four thylacines were captured alive professionals, and the oral-history records of farmers on Woolnorth: a juvenile was snared in August 1924, an and bushmen with their recollections of the disease, adult pair was captured in July 1925, and the last known are data on thylacines killed, both from the Van thylacine definitely recorded from Woolnorth, an adult Diemen’s Land Company Woolnorth property in female, was captured in October 1925. north-western Tasmania, and the Tasmanian-wide government bounty scheme (Destruction of Native Table 1. Thylacines killed or captured alive at Woolnorth, Tigers) raised against the species. 1874 – 1933. Killed or Killed or Year Year The Background Extermination captured captured Rate 1874 7 1894 4 1875 11 1895 6 The data on killed and captured thylacines at the Woolnorth property of the Van Diemen’s Land Company 1876 5 1896 3 (Table 1 - based on Guiler, 1961, tables II and IV; and 1877 8 1897 4 Guiler, 1985, tables 2.1 and 2.3; with minor corrections 1878 7 1898 2 and additions relating to ten specimens from the 1879 1 1899 3 present author’s archival research) suggests a species in decline. Of the 161 thylacines known to have been 1880 5 1900 19 killed or captured on Woolnorth during the 60 years 1881 9 1901 9 from 1874 (the date of the earliest preserved station 1882 4 1902 3 diary and account book) to 1933, a little under half of 1883 9 1903 2 them, some 66 specimens, were killed in the first ten years. In a typical illustration of local species decline, 1884 1904 in the second ten-year period, from 1884 - 1893, only 1885 1905 29 thylacines were killed. Such classic illustration of 1886 1 1906 1 decline did not continue. 1887 3 1907 Significant behavioural change occurred in either the 1888 3 1908 thylacine or human population at Woolnorth towards 1889 2 1909 6 the end of the next ten-year period. In just two years, 1900 and 1901, 28 thylacines were destroyed 1890 6 on Woolnorth. In total, 55 thylacine specimens were 1891 7 1924 1 killed during this third ten year period, from 1894 - 1892 2 1925 3 1903. During this time, no determinable behavioural 1893 5 changes have been recorded for the humans with Australian 2012 Zoologist volume 36 (1) 77 Paddle The background extermination rate at Woolnorth, from then, for a brief period commencing in October 1896, five 1874 to 1899, with consistently reducing numbers of adult thylacines were presented for bounty payment in thylacines killed, by itself was likely to lead to local just six months; such increase reflecting a contemporary extinction. Against this steady decline the sudden, increased ease of obtaining distressed, diseased or dead thylacines ease of capture of thylacines at Woolnorth at the turn of the encountered in the bush. But that was it. No further twentieth century is most likely due to the arrival in north- thylacines from the surrounding Portland District were western Tasmania of the slow and debilitating epidemic presented for bounty payment to the police station during disease as it spread westwards across northern Tasmania. An the remaining twelve years of the bounty’s operation. unidentified bushman, interviewed by Lindsay Crawford at Contemporary newspaper comment also recognized, Long Hill, near Railton, noted a progressive decline of and welcomed, this newfound scarcity of thylacines thylacines in north-western Tasmania. Before they died in north-eastern Tasmania: “Formerly tigers were very off “Tigers were very common around Smithton, & later troublesome … on the North Esk River” (Launceston still, beyond the Arthur River” (Crawford, interview notes Examiner, 22/3/1899). 21-22/11/1952). Harry Wainwright, one of the last “tiger Prior to the spread of the disease, it was extremely unusual men” employed at Woolnorth by the Van Diemen’s Land to find thylacines dead in a snare, and prospective bounty Company, recalled that the disease appeared in north- claimants were officially warned by the Tasmanian Lands western Tasmania around 1899, and was expressed first and Surveys Department of this potential danger: “Tigers of all in thylacines living between the Arthur River and do not choke themselves with the snares, - it is a very Marrawah - Mt. Cameron area, while thylacines living rare thing to find one dead” (Braddon, memorandum to the north of Mt. Cameron initially remained healthy 28/5/1888). But after the appearance of the disease in the (interview 1/10/1972). The manager of the Van Diemen’s mid 1890s, it was another matter. In the wild, diseased Land Company, A.H. McGaw, also referred to the sudden and distressed individuals were easily killed. When snared, disappearance of thylacines at the turn of the century: diseased thylacines tended to stay snared, frequently made “Up to within a few years ago there were a large number little attempt to free themselves, and often these infected of tigers on the Woolnorth Estate” (letter 10/7/1908). Of individuals died as a result of the additional trauma of related interest is the comment by the naturalist, the Rev. capture (Paddle, 2000). Henry Atkinson, who, when passing through Woolnorth in Table 2. Total number of thylacines killed (adults and the early 1900s, noted the absence of Tasmanian devils and young) for each year of the government bounty scheme native cats around Mt. Cameron (Atkinson, 2001, p224). (n = 2,209). A new analysis of the total number of thylacines annually Year Adults killed Young killed Kill Total approved for government bounty payment is presented in Table 2, based on a re-analysis of the Lands and Surveys 1888 58 8 66 Department Ledgers (first undertaken by Eric Guiler (May – Dec) in the early 1960s, Guiler 1961); a careful sifting of the 1889 110 4 114 Lands and Surveys Department correspondence files, 1890 129 2 131 municipal archives, and preserved archival records for the 1891 89 3 92 duration of the bounty from 22 Tasmanian police stations 1892 107 7 114 (Paddle, 2007). (This research is on-going – further police station archives for the 21 years of the bounty remain to 1893 104 8 112 be read by the author – thus the data presented in Table 1894 101 5 106 2, whilst an improvement on previously published bounty 1895 104 5 109 analyses, is not to be taken as definitive.) 1896 122 2 124 State-wide, the bounty shows a similar elevation in the 1897 106 10 116 total numbers killed for the years 1899 to 1902 to that 1898 106 5 111 recorded at Woolnorth, reflecting the arrival of the disease in the less-settled north-west. (Note, however, that the 1899 122 11 133 spread of the disease in southern Tasmania has not yet 1900 138 15 153 been accurately determined.) 1901 157 11 168 Police station records of the bounty from north-eastern 1902 112 18 130 Tasmania reflect the difficulty Wm McGowan began 1903 85 3 88 experiencing after July 1893 in obtaining thylacine 1904 93 17 110 specimens from the north-east for Launceston City Park Zoo. (In fact, McGowan had to await the arrival of the 1905 101 15 116 new century before live thylacines were again obtained 1906 42 1 46 for the zoo from north-eastern Tasmania.) For example, in 1907 47 4 49 the five years, from 1891 to 1895, eight adult thylacines 1908 15 2 17 were presented to, and approved for bounty payment by, 1909 constables at the St. Helens’ Police Station (St. Helens 2 0 2 (Jan – June) Police Station, 1907). No thylacines were presented for bounty payment in the first nine months of 1896. But Bounty Totals 2,050 159 2,209 Australian 78 Zoologist volume 36 (1) 2012 The thylacine’s last straw To the background extermination rate being applied to struck home, the thylacines went down like flies. As fast thylacines because of its pest status, one has to add-in as they died, Melbourne replaced them, such that in the the effects of the disease. The disease increased the ease next 20 months 13 additional thylacine specimens (xxii, of capture, increased the likelihood of encountering dead xxiia, and xxiii to xxxiii) were added to the display. Of these thylacines in the bush, both snared and unsnared, and 19 specimens present or purchased between August 1900 thus materially contributed to the number of bounty and March 1902, all but one had died from the disease by “kills” made. This may be read in both the government 1st December 1902. bounty and Woolnorth data, and suggests that the disease Initially, lesions on the skin were interpreted as instances should enter the foreground in a consideration of the of intraspecific aggression. One of the cubs bred in factors leading to the species’ extinction. captivity (xviii) died on 19th August 1900, and was Age-related effects of the disease are also suggested in the recorded as “Eaten by others”. When one of the parents bounty data. Of the 159 juvenile thylacines identifiably died on 24th May 1901, its bloody, damaged skin prompted procured for the government bounty between 1888 and the comment “Apparently killed by its mate” (Royal 1909, a little over two-thirds (107) were presented for Melbourne Zoo, 1931a). payment in just the first ten years of the initial spread of the Thylacines appeared remarkably placid in captivity in disease, from 1896 to 1905 (Table 3). For this contingency the mixed social groupings in which they usually found table χ2 = 7.8751, p < 0.01, suggesting that, in the wild, themselves. Only two instances of intraspecific aggression juveniles were more susceptible to the disease than adults. were ever observed and recorded in the history of the Table 3. Age separation of raw bounty data, for the zoological garden display of thylacines. In July 1889 one first recognised decade of the disease, versus the other of two male thylacines, the sole occupants of a cage (eleven plus) bounty years. at Launceston City Park Zoo, attacked, killed and ate First Disease Decade Other Bounty Years “a considerable portion” of the other (Trot, 1889). At (1896 – 1905) (1888 – 1895, 1906 – 1909) Washington Zoo, late in the morning of 6th October 1905, a fight, unrelated to any feeding issue, broke out between an Adults 1,142 908 unrelated and isolated pair of thylacines, some nine weeks Juveniles 107 52 after they were first introduced to each other. The male had, the previous year, been successfully housed, without The maximum government pay-out in any one year was expressing aggressive intent, in an isolated pair with a for 168 thylacines in 1901, representing a bounty paid, on mature, previously pair-bonded and mated-in-the-wild average, close to once every two days. Such disease-assisted female (not unsurprisingly, this pairing was without issue). capture of Tasmania’s dominant indigenous predator The new female was one of her offspring, an immature, represented an unsustainable rate of kill likely to lead to the virgin, three-year-old cub, who had “half of one ear bitten rapid extinction of the species. Just eight years later, when off & a bad cut on her head” (Blackburne, 7/10/1905). the bounty was finally terminated in 1909, the government After immediate separation, the pair was later reunited, was paying a bounty on a thylacine, on average, once every and spent a further 48 months living together as an isolated five months. Extinction duly followed.4 couple before the death of the male, without further recorded aggressive interaction, but also without breeding. The Symptoms of the Disease in The post-mortem comments from Melbourne Zoo, above, Captivity on the first two deaths, unrelated to any recorded The first appearance of the disease in zoological gardens intraspecific aggression in the daily record book (Royal around the turn of the twentieth century, was devastating Melbourne Zoo, 1915), were made in the absence of in terms of marsupi-carnivore mortality. While not all any knowledge of the existence of a disease. Certainly, a thylacines (or Tasmanian devils for that matter) died at thylacine body exhibiting significant hair loss and bloody first contact, the episodic nature of the disease eventually skin lesions presents prima facie evidence of intraspecific wore them down, and long-term survivors were rare. aggression. But as the reality of the disease hit home, symptoms of infection becoming obvious in the specimens, Melbourne Zoo and lesions and hair loss appearing on their bodies in the At the beginning of August 1900, Melbourne Zoo had absence of any observed fighting, intraspecific aggression six thylacines on display, an adult male and female pair was ruled out of the equation. Some additional symptoms and their four cubs (specimens xvi to xxi), the product of dying thylacines are noted in the death book: xxiv (a of the only successful breeding of the species in captivity female cub) “Died from Cold” (24/8/1901) – a comment (Paddle, 2000). The vector or specimen that introduced indicative of infection, as distinct from the winterly “Died the disease to Melbourne is unknown. Prime candidates, of cold” – and the juvenile male (xxxii) that died on 9th however, would be the three Tasmanian devils arriving at April 1902 exhibited “sore feet diarrhoea & weakness” the zoo on 16th June 1900. When the disease surfaced and (Royal Melbourne Zoo, 1931a). 4 The few thylacines surviving into the twentieth century, laid waste by human predation and disease, tended to show abnormal, stressed behaviour in their hunting practices, and frequently solitary existence. Oral history records on the behaviour of the species obtained from trappers and old- timers with personal knowledge of the thylacine in the twentieth century, reflect the reality of thylacine existence at the time, but tend to be atypical. Nineteenth-century records of the species – from both Indigenous and invasive European perspectives – identify the thylacine’s typical social and hunting behaviours as based around a small, family group (as summarised in the second paragraph of the Introduction). Australian 2012 Zoologist volume 36 (1) 79 Paddle So rapid and constant proved the deaths, that causes and To two existing nineteenth-century native cats, Melbourne symptoms went unremarked in the death book, to the point added three further specimens between 1st June 1900 and where eventually the deaths themselves were recorded 28th June 1901. All five died between 26th March 1900 only spasmodically. For example, records from the National and 1st September 1901. To the one tiger cat already on Museum of Victoria identify four different dates in July display, Melbourne added six more specimens between and August 1901 on which dead thylacines were received 20th December 1901 and 12th September 1902. Six of from Melbourne Zoo (National Museum of Victoria, 1903, these died in the twelve months between 28th August 1915, 1923), but only one of these departures is recorded 1902 and 26th August 1903. The seventh survived and in the zoo’s death book (Royal Melbourne Zoo, 1931a). respectably lived on in captivity until 23rd February 1905. Furthermore, the bodies of two of these four thylacines To one existing nineteenth-century devil, Melbourne were in such poor condition that no attempt was made by obtained nine additional specimens between 16th June museum staff to save the specimens. No date of death has 1900 and 28th October 1903. Five of these were used in been recorded for the single surviving specimen alive on specimen exchange with other zoological gardens. The 1st December 1902. Later that month another thylacine five remaining incumbents had all died by 5th January (xxxiv) was obtained from northern Tasmania, and this 1904, with the disease confirmed in the Death Book specimen, either because it was now the sole representative, (Royal Melbourne Zoo, 1931a) as being present in the devil display. Melbourne was unable to obtain any further or because it was judged to be the better in physical devils for the next two-and-a-half years, but then they condition, was sent to Antwerp Zoological Gardens in became readily available once again, most, however, January 1903. Unfortunately, though not surprisingly, it not being in the peak of physical condition. In similar appears to have died on board ship before arrival. While fashion to the earlier treatment of the thylacine display, the disease in Tasmania contributed to the ease of bounty Melbourne continuously replaced the devils as deaths claiming, it made trade in live thylacines a much more occurred in the collection. In a little over a year, between difficult proposition. After the arrival of the December 6th May 1907 and 25th July 1908, Melbourne obtained 1902 specimen it was 18 months before Melbourne was eleven devil specimens, two of which were successfully again able to obtain another thylacine. sent to the Zoological Society of London, but the other For the record, like the thylacine, Tasmanian devil deaths nine all died between 17th October 1907 and 2nd May in Melbourne Zoo around this time were initially ascribed 1909. The problem continued, but at a reduced level of to intraspecific aggression (3/5/1899) – and devils being morbidity. Some devils began to survive repeated contact devils this may well have been the case in some instances – with the disease and live to respectable ages in captivity, but certainly for later deaths, after the devil arrivals in June while others continued to succumb to its effects. The 1900, the problem was recognized as disease, described annual report on autopsies of Melbourne Zoo specimens as “mange” (4/3/1902: Royal Melbourne Zoo, 1931a). carried out by the Veterinary School of the University Similar explanations of skin lesions as traumatic wounds, of Melbourne, identified the cause of death of a devil rather than dermatopathies, are recorded for Dasyurids that died in 1911, as due to the “no. lesions”. The only at this time at London Zoo. Up to 1932, Dasyurid skin other Dasyurid autopsied, an eastern native cat, died of lesions were most frequently designated in London as the multifariously-origined “trauma”, together with an “killed by companions” (Canfield & Cunningham, 1993, “impacted rectum” (Stapley, letter 4/3/1912).5 p163). Such explanatory bias, as these authors point out, Launceston Zoo “may have been identified, with histologic examination, as infectious or neoplastic in origin” (p164). Wm. McGowan took over as Superintendent of Before leaving Melbourne Zoo, having addressed the Launceston City Park Zoo in June 1882, and obtained his effects of the disease on its thylacines, a few reflections on first thylacines for display in 1885. Remarkably, the first the effects of the disease in the other marsupi-carnivores occasion in the history of the zoo where expenditure was on display are offered. The precision in recording details of granted for the purchase of stock was not until 1921, when the smaller carnivores at Melbourne Zoo frequently lacks Launceston City Council approved the expenditure of the specificity (and resulting individuation) accorded the £24/9/10 on the “Purchase of Birds” (City of Launceston, larger carnivores. Certainly, some of the smaller marsupi- 1921). Up to then McGowan developed the zoo on the carnivores originated from Tasmania, others from Victoria basis of donations and local exchanges with Tasmanian and the other mainland states, but for many the original animal collectors; offering, for example, Australian locality is unknown. For comparative purposes, native mainland birds in exchange for thylacines (M. Turner, cat, or eastern quolls, Dayurus viverrinus, and tiger cats, or interview 25/10/1992) until, after the sale of specimens spotted-tailed quolls, D. maculatus, are known to live up to to mainland zoos, he was able to build up a small cash three years in the wild and, with the improved knowledge reserve for the purchase of exotic animals for Launceston. and care available in modern-day institutions, up to six Fiscal requirements eventually forced McGowan to start years in captivity. For Tasmanian devils – pre the arrival of negotiations over the sale or exchange of all his thylacines the facial tumour disease – life expectancy was up to six with other zoos. The presence of the epidemic disease years in the wild, and eight years in captivity (Holz, 2008). saw him advertise widely for thylacine specimens (Hobart 5 Melbourne’s only thylacine living in 1911, an adult of unknown sex (xxxvii), died in December 1913. The cause of death was not recorded, but the body was sent to the Veterinary School of the University of Melbourne, for post-mortem analysis and dissection. Unfortunately, no record of the post- mortem has been preserved in the archives of either institution. Australian 80 Zoologist volume 36 (1) 2012 The thylacine’s last straw Mercury, 16/7/1908, 18/7/1908), and necessarily obtain Wolf acts weak & indisposed. Took some chicken & milk” them from further afield than the depleted population (Ditmars, 29/12/1902). The invalid food was apparently in north-eastern Tasmania. It also encouraged him to a success, as it was noted on the following day that the speed up the process of sale and exchange. Between June “Tasmanian Wolf fed well today” (Ditmars, 30/12/1902). 1901 and February 1906 McGowan obtained 24 thylacine This, the first of three bouts of illness, passed relatively specimens (xxvi to xl, xla, xli to xlviii), 16 of which he quickly and without amplified comment. rapidly dispatched to Melbourne, Sydney (Moore Park) The thylacine became ill again for the second time on 2nd and Washington Zoos. Seven thylacines, however, died February 1903 when, under “Illness worthy of note” in within days of their arrival in Launceston Zoo, before any the mammal department’s “Daily Report of Occurrences” sale or exchange could be effected. it was noted, in similar fashion to the Melbourne Zoo records, that the “Tasmanian wolf has very sore feet” Adelaide Zoo (Ditmars, 2/2/1903). This difficulty and pain in movement Adelaide Zoo’s thylacine display had temporarily ceased at was again referred to the next day. The following day all the end of 1896, with the death of specimen xiii that had was apparently well with the thylacine, and it was noted been on exhibit since August 1892. Adelaide managed to that, apart from a lemur which had died and an ocelot obtain nine thylacine specimens between January 1897 killed by some pumas, that “Everything else in good and September 1902, including an adult male (xvii) from shape” (Ditmars, 4/2/1903). Once again, the illness passed Launceston Zoo on 15th June 1898 (Royal Zoological fairly rapidly. Society of South Australia, 1929). Shortly after this The next three and a half months passed uneventfully, but specimen arrived the thylacine display began to wind itself then the thylacine became ill for the third time. On 22nd down. Two specimens died between July and November May it was recorded: “Tasmanian Wolf very lame. Fed well 1898, and two more in January and February 1899 (Hale, last night, taking a pan of milk & all his meat” (Ditmars, 1956). Two thylacines died at unspecified times in 1900. 22/5/1903). The problem continued: “Tasmanian wolf Two more deaths occurred in 1901, in July and September very lame” (Ditmars, 23/5/1903); and the extent of the (South Australian Museum, 1907), and the last thylacine problem was mentioned the next day: “Tasmanian wolf exhibited in Adelaide (xxii) died on 13th September has very sore feet. Den covered with blood” (Ditmars, 1902. No veterinary records or post-mortem comments 24/5/1903). Fortunately on the 25th it was noted have been preserved in the zoo archives for these nine “Tasmanian Wolf began feeding tonight. Took a young specimens. The zoo destroyed the skin and post-cranial chicken” (Ditmars, 25/5/1903) and two days later it was skeletons of two of these specimens, and not even the recorded in the mammal department that the “General skull was preserved from a third death. Six specimens condition of the animals very good” (Ditmars, 27/5/1903). were considered at the zoo to be probably in good-enough After three bouts of this disease, each increasing in condition to be donated to the South Australian Museum, magnitude, the thylacine never experienced the problem and it is possible to specifically identify five of these in the again, and survived in apparently healthy condition until Museum’s current collection. While the skeletal material 15th August 1908, when it was recorded “Tasmanian Wolf was preserved from all five specimens, only one of the sick”, and two days later, on 17th August “Tasmanian Wolf skins was deemed worthy of preservation by museum staff. died” (Blair, 15/8/1908, 17/8/1908). The loss of the last nine thylacines on display, between Beaumaris Zoo July 1898 and September 1902 matches the record of the epidemic disease on the ground in Tasmania, and that For the first fifteen years after the disease became apparent being experienced in Melbourne and Launceston Zoos. in north-eastern Tasmanian marsupi-carnivores in the The known preservation of only one skin from these last mid 1890s, it appeared particularly virulent and thylacines nine specimens is most unusual, and strongly suggestive of particularly vulnerable. During the next fifteen years, their presentation with significant pelagic damage (sensu specimens killed or captured in the wild often exhibited Dumpty, cited in Carroll, 1872). skin lesions, and the trauma of capture, or the addition of being sent to a zoo, could still lead to death. Nevertheless, New York Zoo captive specimens now more frequently emulated the Despite the virulence of the disease at its first appearance, behaviour of the 1902 New York male, in that they would the occasional adult thylacine survived repeated bouts of exhibit symptoms of the disease for a time, but would then the disease and lived into old age. recover. The presence of the disease in Mary Roberts’ Beaumaris Zoo, Hobart, is offered as indicative of these New York’s first thylacine, “a fine male specimen” times.6 (Hornaday, 1903, p59) arrived at the Zoo on 17th December 1902, courtesy of Carl Hagenbeck, the Hamburg animal A female thylacine purchased by Roberts was dead on its dealer. Despite its arrival in good condition, it was noted arrival at the zoo on 6th June 1911, and one of the two on 29th December that the thylacine was ill: “Tasmanian devils that accompanied it died the following day. The 6 In this paper, the name “Beaumaris Zoo” has been reserved for just the privately run Beaumaris Zoological Garden, established by Mary Roberts at Sandy Bay, on the then outskirts of Hobart, with its orientation towards the display of Tasmanian and Australian species. After Roberts’ death in 1921, her collection formed the basis of the short-lived (1922 – 1937) Hobart City Council’s zoological garden at the Domain, which officially preserved the name Beaumaris in its title, but now with a changed orientation towards the display of exotic species. This second iteration of Beaumaris Zoo is designated herein as “Hobart Zoo”. Australian 2012 Zoologist volume 36 (1) 81 Paddle thylacine’s body was kept at the Zoo for some little time, a suspiciously short and troublesome period of display while Roberts enquired whether Prof T.T. Flynn at the in New York, some 17 months after first-known contact University of Tasmania, was interested in preserving its with the disease. internal organs. Among the marsupi-carnivores then on Whilst on the topic of Beaumaris Zoo, a few observations display in the zoo, to serve as potential carriers for the on the marsupi-carnivore disease in Mary Roberts’ devils vector of the disease, were at least six Tasmanian devils are offered. Mary was certainly enamoured with her and an adult female thylacine (iii) and her male cub (vi), devils, became the first person to breed them in captivity both captured at Woolnorth in 1909. (Roberts, 1915), and, all told, between April 1908 and A large, adult male in fine condition (viii), with an August 1920 kept at least 82 specimens in captivity.7 unblemished skin, then arrived at the Zoo from Tyenna There is no mention or indication of disease amongst on 12th August 1911 (Paddle, 2008). Later in the month, the first 21 Sarcophilus specimens displayed in the zoo, this adult male developed a lesion on his tail. Concerned between April 1908 and April 1911. The dates of death, about his symptoms the veterinarian was called for: or last reference to continued existence within a collection “Mr Ritchie Veterinary surgeon came to see the spot on are known for sixteen of these specimens. None of these tigers tail, sent powders and medicine” (Roberts, diary sixteen survived for less than five months in captivity, and 25/8/1911). The male recovered, but the adult female one of the pouch young obtained in October 1908 (Sviii) then fell ill some weeks later. No specific symptoms were died in Melbourne Zoo in March 1914, after 65 months recorded on this occasion, merely a note that “Had the of captive existence. Then three consecutive devils (Sxxii Veterinary Surgeon to see the female tiger in the morning, to Sxxiv), obtained by Roberts between 15/5/1911 and he thought she had a severe cold” (diary 11/9/1911). She 6/6/1911, from three different localities (Triabunna, Little also recovered, and did not trouble the veterinarian again. Swanport and the Lakes District) were all dead within Her specific date of death is unrecorded, falling in a brief 24 hours of their arrival. For the first time, in 1911, Mary hiatus in Roberts’ record-keeping, but appears to have began to refuse the offer of devils specimens, particularly taken place around March 1913. The adult male died those from Triabunna – even refusing the tempting offer of on 9th March 1915. Writing later about the demise of the a mother with pouch young caught by E.E. Ford (Roberts, Woolnorth female and Tyenna male Roberts noted: “The diary 28/8/1911, 29/8/1911). Until 1912 Mary paid the pair were both old & died from that cause eventually” devil trappers on the day, if they arrived with their catch (Roberts, letter 27/3/1919). in person, or within one or two days of an unaccompanied devil’s arrival, via postal note. But in 1912 she began to The remaining young Beaumaris male (vi), whilst he delay immediate payment for devil specimens, sometimes remained in Roberts’ care, showed no sign of the disease for more than three months (in the case of specimens Sxli prior to his departure for London Zoo on 28th September and Sxlii), if she found them diseased on arrival (Roberts, 1911, just 17 days after his mother required veterinary diary 26/9/1912, account book 30/12/1912); and after attention. He arrived in London (as specimen xix) on 21st April 1912 she refused to purchase any further devil November 1911, but did not join the other thylacines specimens on offer from the trappers Ford and Harry on display, one of which was his male sibling. He was Smith of Triabunna. placed in the Zoo’s sanatorium and remained there for over seven weeks (a suspiciously long quarantine London Zoo time), during which time his sale to New York Zoo was For the record, the three thylacines housed nearby to negotiated. He left London Zoo on 10th January, and the quarantined Beaumaris specimen in London Zoo arrived at New York Zoo (as specimen ii) on 26th January showed no obvious external sign of the disease on their 1912. Hornaday wrote to Roberts (letter 6/4/1912): “We bodies. One of them became ill, and was daily recorded have at last secured a fine Tasmanian Tiger, which is as “unwell” between 30th January and 18th March 1913 living in our Small-Mammal House, in good condition, (Zoological Society of London, 1913), but it recovered and seems to be enjoying life”. But despite this promising fully and lived for at least a further 15 months. All three start, after only ten months of American display, he died London thylacines, however, died in 1914. The first (xvi) on 20th November 1912. Unfortunately, the daily reports on 5th June from “Chronic Nephritis”, another (xvii) on 20th for the mammal department and veterinarian’s office November from “Peritonitis”, and the last (xviii) on 25th for 1912 have not been preserved in the New York Zoo December, also from “peritonitis”, as a result of “perforation archives. The only comment about its departure was of colon” (Zoological Society of London, 1916). Of the that “while it arrived in good health, it was so nervous eleven thylacines that died in London Zoo between 1884 and unreconciled to captivity that it lived only a few and 1931, peritonitis was the identified cause in five cases, months” (Hornaday, 1912, p71). This comment is rather enteritis and nephritis accounted for two further deaths, anomalous, as the specimen had certainly proved well- and four departed without exciting post-mortem comment. adjusted to captivity in Hobart, having been caught, Peritonitis, due to gastric ulceration, is a common marsupial along with its mother and two of its siblings, as a young stress-related condition as a response to prevailing captive cub, around ten months of age, between late May and conditions (Canfield & Cunningham, 1993). Similarly, early June 1909. It died at a relatively young age, after nephritis was a common necroscopic diagnosis across all 7 The total of 82 displayed is a minimum estimate, and makes the assumption that all four devils that escaped during the night of 7/5/1911 were recaptured, and later sold or exchanged. It is unusual, however, that such hypothesized success in recovery failed to be recorded in Roberts’ diary. Australian 82 Zoologist volume 36 (1) 2012 The thylacine’s last straw marsupial groups displayed in London Zoo (Canfield & Hobart Zoo Cunningham, 1993; Crisp, 1860; Plimmer, 1915). The No zoological garden proved able to obtain thylacine female thylacine that died on 5th June 1914 of “Chronic specimens directly from the wild in 1926 or 1927. At Nephritis” is of greater interest here, howsoever, given the Hobart Zoo, under the curatorship of Arthur Reid (and later veterinary analysis of nephritis accompanying deaths later his daughter Alison), come the end of 1927, two specifically ascribed to the epidemic disease in thylacines female thylacines remained on display. But, for a brief at Hobart Zoo. time in 1928, thylacine availability changed, for the The marsupi-carnivore disease, however, certainly made worst of all possible reasons, as a particularly virulent inroads on London’s devil collection. The specific life- strain of the disease appeared in the species once again, histories are determinable for all five Tasmanian devils on and distressed thylacines appear to have become more display in London Zoo at the beginning of January 1911, readily encountered and captured in the bush. To the two prior to the unprecedented incidence of ante-mortem already in stock, a further seven specimens (xx to xxvi) comment found in the Daily Occurrence Records (Zoological were added in the first four months of 1928. By the end Society of London, 1911a, 1912) that preceded their deaths of the year, however, only two of these nine specimens from the disease. The oldest, specimens Sxv and Sxvi, remained alive. While such virulence hardly represents were both males that had arrived from Melbourne Zoo effective use of a host species by a disease organism, it is in June 1908. Specimen Sxix, of unrecorded sex, arrived readily explicable, given that the disease does not appear from Beaumaris Zoo, also in June 1908 (together with two to have been restricted to a single host species (de Castro companions sold/exchanged to other institutions). The last & Bolker, 2005; McCallum & Dobson, 1995). two specimens, Sxx and Sxxi, of unrecorded sex, arrived Arthur Reid considered that the disease was “perhaps from Beaumaris Zoo in November 1908. The source of distemper that they had caught from the trapper’s dogs”, infection for London’s devils is unknown. The disease was but Alison Reid was less convinced: “there wasn’t any not recorded at Beaumaris Zoo until May 1911, well after sign of discharge from the eyes and nose like you see the despatch to London of all of specimens Sxvii to Sxxi. with a [distempered] dog … they just died” (Alison While the epidemic disease had appeared in the marsupi- Reid, interview 1980). While it is now known that carnivores at Melbourne Zoo in 1900, the devils received canine distemper virus can spillover to non-canid species in London from Melbourne (Sxv and Sxvi) showed no sign (Roelke-Parker, et al. 1996), against the direct spillover of the disease for the relatively long period of time between of distemper between the two species is a contemporary June 1908 and January 1911. suggestion that it was only after the wild dogs in the bush One Tasmanian devil was recorded as “unwell” from 24th to had died of distemper that thylacines became common 26th January 1911. Two Tasmanian devils were recorded as (Tasmanian Mail, 9/8/1884). “unwell” from 28th to 31st January. One of these devils (Sxv) An adult female with two pouch young (xx to xxii) died on 1st February. The second devil remained “unwell” entered the zoo on 10th January 1928. A product of on the 1st and 2nd February, joined by a third “unwell” devil the previous peak breeding season in early spring, the from 3rd to 5th February, with a fourth devil also recorded as cubs “were fairly advanced, they were just about on the “unwell” from 6th to 7th February. On 8th February Sxx died point of emerging when they came in” (Alison Reid, and was autopsied. As in previous zoological garden records interview 27/2/1992). The new arrivals joined the two for the first incidence of the disease, the lesions associated resident females in the thylacine cage on the hill at the with the disease were described as “Injury, suppuration back of the zoo. of wound” together with “gastric ulceration, tapeworms” (Zoological Society of London, 1911b). Specimens Sxvi A fortnight later, on 24th January another thylacine and Sxix recovered, but Sxxi remained “unwell” on 8th and (xxiii), a juvenile male arrived at the zoo, brought in 9th February, before dying on 10th February. Post-mortem some haste by its captor, T. Hunt of Tyenna. Obviously comment again interpreted the lesions as “Injury”, together ill on its arrival - “There was one very sick one that with “dilated heart, nutmeg liver, congestion of lungs” came in with a virus, it came from Tyenna” (Alison (Zoological Society of London, 1911b). On the same day, Reid, interview 20/10/1995) - it was not added to one of the two remaining devils was recorded as “unwell”, the collected thylacine display, but caged alone near and remained so until 14th February. Both devils were then the front entrance of the zoo. The photographer Ben recorded as “unwell” from 16th to 19th February, but one Shepherd, of Sandy Bay, visited the zoo within days of these recovered. A devil was recorded as “unwell” on a of its arrival and took a glass half-plate negative of daily basis for the next 53 days, between 20th February to the diseased specimen (Figure 1). Shepherd provided 13th April, then both appeared healthy and failed to trouble the scientific illustrator and photographer, Norman the daily records for some little time. Four months passed Laird, with a copy from the negative, and details of before the disease returned again. One devil was recorded the specimen and its photography. As recorded in an as “unwell” from 26 August 1911 to 29th September, both unpublished manuscript by Laird: “The animal died the appeared healthy on 30th September, then the other devil day after it was photographed, and it does not represent became “unwell” from 1st to 19th October. Both successfully the species in good condition” (Laird, ca1978). Despite survived this second bout of disease, but they succumbed the relative isolation of its display, alone in its cage at the when it returned for the third time two months later. Sxvi front of the zoo, “that was where the sick one came but died on 2nd January, and Sxix died on 23rd January 1912. the virus seemed to spread right up to the other [cage]” Neither of these specimens was autopsied. (Alison Reid, interview 20/10/1995). Australian 2012 Zoologist volume 36 (1) 83 Paddle thylacines to die were not offered as museum specimens. Daily, rather than weekly, rubbish collection from the zoo only commenced in 1931 (Reserves Committee, minutes 20/7/1931), and, as dead thylacine bodies were a bit too large to hang around for up to a week, the corpses were buried “in the zoo, up on the bank” (Alison Reid, interview 25/6/1992). The only specimen whose body was deemed worthy of preservation was that of a juvenile female that died on 18th July 1928, was purchased by the Tasmanian Museum and duly preserved as a mounted display. Two thylacines survived the year, although continuing at times to show periodic evidence of the disease. They were both on display when Arthur Reid drew up the “statement of stock” on 31st October 1929, but one of them died the next day: “The Curator … reported that one Tasmanian Wolf had died of kidney disease” Figure 1. The diseased Tyenna male (xxiii), caged separately (Reserves Committee, minutes 1/11/1929). The body in Hobart Zoo, late January/early February 1928, on the was sold to MacKenzie for anatomical purposes. The penultimate day of its existence. (Shepherd) designation of “kidney disease” is interesting, and has This juvenile Tyenna male died at the end of January, parallels with the thylacine death at London Zoo on or very early into February 1928. No further thylacine 5th June 1914. Alison Reid was persistent (interview deaths were recorded in the monthly stock mortality list 24/6/1996) in her opinion that all thylacines present from 14th February to 13th March. In fact, an additional in 1928 eventually died of the epidemic disease. The juvenile female thylacine (xxiv) had arrived on 28th veterinary surgeon, Dr C.M. Sprent visited the zoo February. It was not until two months after the young twice, in early October and again in late October or Tyenna male had arrived (and rapidly departed), that early November 1929 to examine the thylacines (Alison the animals displayed in the thylacine cage proper, up Reid, interview 1980; Reserves Committee, minutes the hill at the back of the zoo, started dying. As Alison 14/10/1929, 18/11/1929), and is the likely source for the Reid recalled: inference of renal failure. • they just sort of keeled over and died. (interview The last of the zoo’s thylacines present in 1928 died of 27/2/1992) the disease five months later, missing from the stock report of 31st March 1930. “The Curator reported • We were pretty badly hit, because all of ours died, one that the Tasmanian Wolf had died of kidney disease” after the other, they all died. (interview 24/6/1996) (Reserves Committee minutes, 14/4/1930). Presumably The first to die, on 20th March 1928, was one of the newly the identification, once again, of death from “kidney arrived cubs, a female (xxi). Its skin was in good condition, disease” was a diagnosis of the veterinary surgeon’s, but it and the cub was sold to the Tasmanian Museum, where it is not known whether Sprent was called to the zoo at the was mounted by Alison Reid. Then, on 27th March Arthur time of the thylacine’s death, for the record of accounts Reid reported to the Reserves Committee meeting that paid between 18th and 30th March 1930 are missing from “one of the female Tasmanian Wolves had died”, not the the zoo archives. It was seven months before the zoo was 10th January mother, but one of the two females on hand able to obtain another thylacine for display. at the start of the year. The skin of this specimen was in poor condition. Although externally unsuitable for public It is not known for certain whether the Hobart Zoo loss of display, her body was sold for internal display to Colin all nine thylacines to disease, between January 1928 and MacKenzie of the Museum of Comparative Anatomy, March 1930, was paralleled elsewhere. James Harrison, Melbourne (the collection later moving interstate to estate agent and animal dealer in Wynyard, recorded become the Institute of Anatomy, Canberra). the purchase of two live thylacines from an unidentified source around April 1928: a juvenile for £7/10/- and an With three thylacines lost already this year, and with their adult for £20. Both were sold to Colin MacKenzie in May value as exchange or sale specimens appreciating, the 1928. The juvenile had died in Harrison’s care, and its Reserves Committee promptly sent Arthur Reid to Port body was sold to MacKenzie for just £12, but Harrison Davey, as he “had been informed there was a number of demanded and received £50 for the live adult (Harrison, Tasmanian Wolves there” (Reserves Committee minutes, 1931) which was immediately deposited in Melbourne 27/3/1928). The visit was successful, Arthur Reid returning Zoo. Melbourne had two thylacines on display in 1927, with a pair of thylacines (xxvi and xxvii) on 17th April 1928. and obtained just two more specimens from the wild, on From April 1928 onwards, thylacine deaths ceased to 11th May 1928 and 16th October 1929. Unfortunately, be significant events, becoming almost matter-of-fact due to the loss of archival records from the Zoo, as well occurrences, Arthur just “used to say ‘another tiger died’” as the Institute of Anatomy (the repository of the bodies (Alison Reid, interview 27/2/1992). Because of the poor of the last six Melbourne Zoo thylacines on display), quality of the skins, possessing significant areas of active the death date of only one of these last four Melbourne lesions and hair loss; the bodies of three out of the next four specimens is known, that of the last thylacine received, Australian 84 Zoologist volume 36 (1) 2012