Thylacines and Dingoes

By Branden Holmes

Early European maritime visitors to Australia recorded the close relationship between the aboriginal people and wolf-like dogs. The first known mention of dingoes appears to be that of Jan Carstenszoon in 1623 around Queensland's Cape Yorke Peninsula. Later early encounters include a "yellow Dog" near Jurien, Western Australia in 1697 (Abbott, 2008), William Dampier's crew (1699), the HMS Endeavour (1770), and the First Fleet (1788). An illustration of the "dog of New South Wales" was made by Arthur Phillip, first colonial governor of NSW, not long after the First Fleet landed in 1788. It was this illustration that was the basis, first of Robert Kerr's now abandoned name for the dingo Canis antarticus (Kerr, 1792; but see ICZN, 1957), and then of today's accepted name Canis dingo given by Friedrich Meyer in 1793. Though today more often referred to as Canis lupus dingo or Canis familiaris dingo, in order to emphasise its domestic origins.

Early European encounters with the thylacine on the other hand are much less clear. Paddle (2000:3) suggests a similar timeframe, the seventeenth and eighteenth centuries, for the first European encounters. The account of Abel Tasman's crew finding "tyger" footprints on Van Diemen's Land (now Tasmania) in 1642 (Rembrantse, 1682), is often reproduced. However, the footprints of a thylacine are decidedly more dog-like, and thus not likely to instill fear in those who come across them. The cute and cuddly wombat, however, has relatively long intractable claws that are likely to leave in fear those who are not familiar with the benign possesser of such fearsome hardware (Guiler & Godard, 1998; Mooney, 2014). It would be easy to mistake a complete print for that of a ferocious tiger-like creature. Thus the description of "tygers" in Tasmania, and the thylacine often being referred to as a tiger due to its stripes, is probably pure coincidence.

The safest of Paddle's suggested pre-1805 records of European contact with thylacines is that on 13 May 1792, by members of the French La Récherche expedition. Made more secure by a re-translation of the relevant phrase by Neil Murray (Paddle, 2000:3). While the earliest two post-1804 encounters seem to be those on 30 March 1805 (Anonymous, 1805) and 2 May of the same year (Nicholls, 1977; journal entry for 18 June). With a smattering of thylacine encounters over the first decade of settlement. Doubtless a product of both the small settler population and the relative rarity of top predators compared with their prey. In his scientific description of the species (read in London, 21 April 1807), George Harris mentions that only two specimens, both males, had been taken since settlement in 1803 (Harris, 1808). One being Paterson's specimen killed on the hill, an average of one every two years. A rate that would sadly climb and climb in years to come. Indeed it would appear to have taken 17 years before a specimen was taken alive (Anonymous, 1820).

The dingo din' go to Tasmania?

There is no evidence to suggest that the dingo ever occurred in Tasmania (contra Krefft, 1868; see Allport, 1868; Gill, 1953:89; Jones, 1968). Either it never arrived, or it died out. The adaptibility of the species is evident on the mainland, where it occupied every single habitat from the deserts to the snow-laden Australia Alps, until it was exterminated from much of its former range (Corbett, 1995a). A range greater even than that of the red fox (Vulpes vulpes), which even in good years does not extend to the top of the continent (Coman, 1995). There is therefore no good reason to suppose that Tasmania was ecologically closed off to the dingo. Moreover, if it did arrive it would undoubtedly have a close relationship with the aboriginals like European-introduced dogs later did (Anonymous, 1826), somewhat contrary to the situation on the mainland (A New Correspondent, 1835). Beyond the obvious logistical fact that it couldn't get there on its own, the relationship would have been mutually beneficial (e.g. hunting, companionship, security etc.¹). A situation mirrored all over the Pacific, where now extinct breeds of dog thrived alongside their Polynesian seafaring owners on many Pacific islands (Williams et al., 2018). Thus creating an extra buffer against extinction.

So it appears that the dingo never reached Tasmania. On the face of it this is a curious fact. The dingoes ancestors probably came from South-East Asia through New Guinea and across the Torres Strait. The latter is about 150km wide at its narrowest, while the Bass Strait which separates Tasmania from the mainland is just under 200km at its narrowest. An extra 50km is certainly a substantial difference, even for skilled seafarers. However, both straits are conspicuously traversable by utilising islands dotted along the way. In the case of the Bass Strait, through the Kent and Furneaux Groups of islands. Thus neither journey would necessitate a strictly overwater affair, and hence each could avoid at least some of the dangers thereof. Thus geography alone cannot account for why the dingo never arrived in Tasmania.

The only conceivable major difference between the two straits is the sailing conditions. The famous Roaring Forties, which produce howling westerly winds in the region of the forties latitude. As well as the sheer ferocity of the waves at times. Which combined with gale-force winds and freezing temperatures (even above the water) made the 1998 Sydney to Hobart Yacht Race the deadliest of all time. Six sailors lost their lives as the 115-strong fleet headed into what can only be described as hell on Earth. Even with all of the modern technologies at their disposal, only 44 yachts crossed the finish line that year. While it is certainly possible to play up the dangerousness of the Bass Strait, evidenced by the rather mundane journey of the Spirit of Tasmania most days, such a ferry is huge and thus not as amenable to the influence of the ocean as much smaller vessels are. Its shallowness presents untold dangers, which aboriginal sailors and fisherman would have been well aware of.

Given the unfavourable sailing conditions, there are three basic hypotheses to consider. Any one of which in isolation could in theory explain the lack of contact between mainland and Tasmanian aboriginals. Although the real answer probably lies in a combination of them: a) there was no great need to sail far from the shore, since fish, turtles, dugong etc. would have been more than plentiful close to the coast, b) some kind of taboo against sailing in the direction of Tasmania, whether or not the people were aware of that distant land, and c) contact would more than likely have been accidental, thus not likely to result in enough dingoes in a short span reaching Tasmania for a viable population to arise. A fourth possibility, in the opposite direction, namely deliberate contact, is what I am arguing against by proxy of the evidence of the absence of the dingo from Tasmania. It is inconceivable that there would have been regular contact without dingoes being involved.

Aboriginal oral tradition, when properly understood and treated with respect, can provide us with evidence for hypotheses that would otherwise be almost impossible to make a case for (Burbidge et al., 1988; Abbott, 2008; Burbidge, 2017). Thus the seeming complete lack of evidence for dingoes in Tasmanian aboriginal oral tradition is evidence that it was at least largely absent from the island. It is of course possible that a tiny number of dingoes were brought to the island and either died out due to inbreeding, were consumed, killed by disease etc. Moreover, Tasmanian aboriginal oral tradition is less completely known than that of mainland aboriginals. While Tasmanian aboriginals survive today, descendents of their full-blood relatives, they have lost an untold wealth of knowledge (their languages have all been lost). Which might have included knowledge of the Australian dingo. So that its absence from known oral tradition is not strictly proof of its absolute absence.

In defence of the hypothesis that dingoes have (largely) been absent from Tasmania, we can also point to the cultural and technological differences between mainland and Tasmanian aboriginals which must have accumulated as a result of their physical isolation from each other. It is well known that Tasmanian aboriginals were more technologically primitive than their mainland counterparts at European contact. For example, they lacked the boomerang even though it would have been of great benefit in the last several thousand years, as the mammalian aspect of their diets increased significantly. A state of affairs that resulted in Victorian minds, gripped by thoughts of evolution, in seeing their demise as "natural". Nevermind the fact that they survived in such a "primitive" state without issue until European contact. Whatever the cause of their relatively primitive state, certain tools would have undoubtedly provided advantages to the Tasmanian aboriginals had they had them. So that their absence is something of a proxy for the absence of mainland influence on their culture.

The final reason to suppose that the dingo never arrived in Tasmania is the ecological results of European dogs being introduced to Tasmania. Feral packs became a problem starting in the late 1820's, resulting in livestock losses increasing dramatically, with the thylacine being falsely blamed. They also displaced thylacines, forcing the latter to retreat to the less inhabited and more mountainous areas of the island (Anonymous, 1884; Boyce, 2006,2008). Before canine distemper took hold and the feral packs slowly died out during the mid nineteenth century (Ibid.). The thylacine population thus began to increase again, albeit with a significantly smaller gene pool. Accounts of thylacine-dog encounters paint the thylacine in a somewhat favourable light, able to fend itself against one or two domestic dogs (Paddle, 2000). Which has led Paddle to conclude that "there is little evidence" that European dogs contributed directly to the thylacine's extinction, contradicting the above.

The evident bias in Paddle's case is that he does not take into account the relevant differences between dogs obeying their masters and a psychologically independent pack of hardened feral dogs. The latter have to be more ferocious and cunning than the former in order to eke out their existence. Moreover, displacement is just as valid a cause of extinction as predation, since both result in a population reduction. There is only so many thylacines that a given area can sustainably support before either fatal thylacine-thylacine competition or starvation starts occurring. Paddle notes that feral packs of dogs tend to arise when aboriginal people have been displaced or murdered. The genocide of the Palawa people certainly would have forced their dogs to adjust to feral life. Thus at least in part explaining the origins of the feral packs that were a serious pest on the Tasmanian sheep industry, and turned some farmers whole-heartedly against thylacines.

Post-European mainland thylacines?

The scientific consensus is that thylacines became extinct on the mainland around 3,200 years ago. Radiocarbon dated specimens vary greatly in age but increase towards a younger date, finally clustering at around the 3.5ka mark before petering out completely (White et al., 2018b). It must be pointed out that some bias is evident in the dataset used to infer an extinction chronology, as the fossil sites from which dates are taken are almost exclusively from southern Australia. Thus there is some potential for northern Australia to have been a refugium for the species, but future dating of northern Australian specimens will either support or disconfirm the currently accepted chronology. It must also be noted that there have been palaeontological claims of more recent specimens (Archer, 1974; McDowell, 1997), but these have been overturned as a result of re-dating (White et al., 2018b).

There was also the extraordinary discovery of the mummified thylacine carcass (nicknamed "Old Hairy") in a Nullarbor cave², by David and Jackie Lowry in October 1966 (Lowry & Lowry, 1967). The state of preservation lead both the discoverers and Atholl Douglas to suppose that its age was less than 2,000 years old, and possibly only several years old (Ibid.; Douglas, 1990). The true age of the specimen, established through radiocarbon dating, was in excess of 3,000 years old (Partridge, 1967; Lowry & Merrilees, 1969; Merrilees, 1971). However, Douglas persisted in his belief that the specimen was very recent, for several reasons, none of which hold up (Williams, 2014). There is also, of course, the many thousands of sighting reports from the mainland (e.g. Healy & Cropper, 1994; Heberle, 2004; Opit, 2014; Smith, 2014).

Although almost ubiquitously known today as the thylacine or Tasmanian tiger, the species had innumerable other names during the nineteenth and twentieth centuries (Guiler & Godard, 1998:15; but see my own list). These include variations on the term 'hyena', so that a very early newspaper report has been hypothesised by The Thylacine Museum (5th revision) to possibly refer to two instances of post-European mainland thylacines:

"Two animals of the hyena kind were seen at Campbell's Island by hunting parties belonging to the Mary and Sally ; from the description given of which they appear to have been of the same species with an animal killed at Port Phillip in 1803." (Anonymous, 1811)

Such an early report does not make one very confident that the thylacine is the only, or even most probable, identification. Especially when one realises that the Campbell's Island referred to is actually the subantarctic island belonging to New Zealand (Crowther, 1932). Nevertheless Robert Paddle has raised the possibility of at least two relict populations of thylacines on the mainland until at least the 1830's and 1840's (Paddle, 2000:22-23). One in South Australia and the other in the Blue Mountains (NSW), as he seems to implicitly dismiss the idea of them constituting a continuous population. His suggestion, however, is based upon a completely flawed case. He cites three lines of evidence: three articles by a naturalist called 'Cambrian' (Cambrian, 1855a,b,c), a lecture by Dr. John Palmer Litchfield (Anonymous, 1840), and aboriginal oral tradition (Tunbridge, 1991). None of these hold up to scrutiny, and worse still, other considerations render his bold claims even more objectionable.

The author Cambrian's true identity is unknown³, and thus the veracity of their writings about the thylacine cannot be ascertained. The significance of their claims means that we need positive evidence, not merely anonymous comments. Given Cambrian's use of a pseudonym, it seems impossible to check their background, thus rendering their comments moot. But Paddle claims that we may infer Cambrian's competence by "a frank apraisal of the accuracy of the non-marsupi-carnivore material...in light of European marsupial knowledge at the time" (Paddle, 2000:23). In reality, Cambrian's competence cannot be gauged by any such method. The juxtaposition of valid scientific observations with bold unverified claims does not render the latter more likely to be true. Especially since we simply do not know the source for their non-marsupi-carnivore material in the articles. And we can't start simply assuming Cambrian's reliability in order to take them at their word as having made the observations themselves.

But let's assume that Cambrian was a competent naturalist and that they did indeed examine two dead thylacines on the mainland. It is far more likely that the actual geographical origin of the specimens was Tasmania rather than the mainland⁴. Whether as a result of an accident or a lie at some point. Instances of wrong locality data being attached to specimens are certainly not unknown. In fact an excellent example of this (see appendix 1) is the claimed recent occurence of the echidna genus Zaglossus in the Kimberley region of Western Australia (Helgen et al., 2012). Historically thought to be confined to New Guinea, a specimen labelled as coming from the Kimberley region was re-discovered in a museum collection. They made a case based upon fossil remains, aboriginal rock art, the living memory of aboriginals, and of course the specimen labelled as originating from Australia (collected by a competent naturalist, John T. Tunney, then director of the Western Australian museum). Conservationist and WA mammal expert Andrew Burbidge evaluated their case and found it to be seriously flawed (see appendix 1), concluding that the "most plausible explanation is that the tag on the specimen came from another animal" (Burbidge, 2017).

Back to Paddle's own case, Cambrian cites both the explorer Charles Sturt and South Australian Governor Sir George Grey as sources relating to mainland thylacines. Unsurprisingly Paddle cannot find any such mention of mainland thylacines in the writings of either (Ibid.). Worse still, Paddle cites a lecture by Dr. John Palmer Litchfield, which claims that a bounty was placed upon thylacines in South Australia. Or at least according to Paddle, who has a penchant for interposing his own interpretation of other people's writings. The "province" referred to is completely ambiguous, but that doesn't stop Paddle from asserting that it is South Australia. If such were true, mainland thylacine research today would look very different. We would find newspaper reports lamenting the devastating effects of thylacines upon livestock, as we do in fact find in Tasmania. So that even if the bounty itself, like some in Tasmania, is scantly attested to (as Paddle himself has uncovered), the motivation for the bounty (i.e. claimed thylacine attacks on livestock) in the first place would be easy to find. Alas that is not the case.

In reality 'Dr.' John Palmer Litchfield was a notorious conman whose writings about the thylacine cannot be relied upon in any serious way⁵. A serial liar and pedagogical fraud. It is very worrying that Paddle is completely unaware of this fact. Since even a cursory look into Litchfield's background would establish that all is not right, such as via an internet search. Granted Paddle did not have access to many of the electronic resources we do almost two decades later, as the internet was still relatively new. But by the same token, his university studies and access to archival material would have more than made up for this, giving him access to information that we members of the public today still do not readily have access to. A background check of the author of a given source should be one of the (if not the first) early tasks of the researcher. Being taken in by a conman is a sign of severe intellectual myopia.

Paddle's last line of evidence is aboriginal oral tradition. The identification of the marrukurli of Flinder's Ranges aboriginal oral tradition with the thylacine is suggested as a possibility by (Tunbridge, 1991:48) because "several Adnyamathanha elders have believed them to be identical". She notes, however, that "there is no certainty" that this is so. Moreover, the marrukurli appears in both the Dreaming and other oral traditions which "suggest the possibility of their bodily existence in the region in the not-too-distant past" (Ibid.). Tunbridge's picture of the marrukurli does not pick out the thylacine as being the obvious match. If it is a single type of animal it is more likely referring to brindle and/or starving dingoes (with their ribs showing), hanging around settlements hoping for a meal. After all there are documented unprovoked fatal attacks on humans by healthy dingoes, while the several alleged instances of thylacines attacking people in Tasmanian are either of sick or injured animals, or it is hard to tell who the actual aggressor was.

The marrukurli comes across as more like a blend of fact and fiction, serving a deeper purpose: warning children not to stray too far from the adults at night. It is said, as is so often the case with large predators, that keeping up a fire wards them off. With the comforts and security of modern living it is easy to forget that Pleistocene people, indeed many people today, must have been so concerned for their safety. Until 50,000 years ago Australia was inhabited by Megalania (Varanus priscus), a giant goanna twice the size of a Komodo dragon. As well as other terrifying predators like the Marsupial lion, whose bite force is pound-for-pound one of the strongest known of any animal, living or extinct. Paddle (2000:23) thus ignores the serious doubt surrounding the identity of the marrukurli, and simply asserts that the two are identical. In summary it is clear that Paddle has unquestioningly pulled together any "fact" that props up his belief that thylacines survived on the mainland until after European colonisation.

A unique problem for Paddle

Paddle's problems are confounded by the idiosyncratic position that he finds himself in. On the one hand he "suggests the possibility" that at least two relict populations of thylacines existed on mainland Australia until at least the 1840's. But on the other hand he claims that they are definitely extinct today, not just in Tasmania and on the mainland, but also on New Guinea. He notes that the species disappears from mainland aboriginal oral tradition during the 1840's and 1850's, corroborated by its absence from contemporary species lists by scientists, which he seems to take as representing the near-terminal extinction chronology of the species (Paddle, 2000:22-23). A few individuals could in theory have lingered on in such low density that not even aboriginals were aware of their continued survival, but that would also lead to a lack of breeding. The overriding problem is that Paddle is largely bereft of a mechanism to explain the species' extinction on the mainland, especially since his inferred extinction chronology is much shorter than that on Tasmania. 

For Paddle, the claim that dogs (whether dingoes on the mainland or European dogs in Tasmania) were a direct cause of the extinction of the thylacine is a symptom of placental chauvinism. The alleged superiority of the placental dingo, with its bigger brain, larger social structure, and broader diet, is a myth. Driven and perpetuated by an innate desire to rid ourselves of the guilt of driving the thylacine to extinction. Their real contribution to the extinction of the species in Tasmania is negligible, while on the mainland they were ancillary to the increased hunting activity that their co-operation with their aboriginal companions allowed. For Paddle then, there can be no valid case made that dogs helped significantly to cause the extinction of the species. In Tasmania it was the false persecution of the thylacine as a sheep killer, which led to several unmitigated bounties, and the appearance of an epizootic disease at the turn of the 20th century (Paddle, 2012), which ultimately led to the species' demise.

Paddle cannot apply either of these extinction causes to the mainland thylacine. The bounty that Paddle mentions can reasonably be said to have never taken place since its only source was a conman, and the epizootic disease that Paddle discusses arrived far too late. Moreover, Paddle needs one or more extinction causes that are even more acute than these two in conjunction. It is safe to say that if it were not for humans (whether directly or indirectly) the species would undoubtedly still exist today. Going back to the year 1800 on the mainland, the thylacine population would almost certainly have been relatively healthy. With no bounty on them, and no good reason to suppose that the population would have been unhealthy⁶, the two populations could have exceeded Tasmania in terms of total population size. But by the 1850's it was almost, if not already, extinct. A period of a mere 50 odd years.

If we take the Tasmanian thylacine situation from the years 1870-1920, encompassing both the 1888-1909 government bounty and the rise and spread of the epizootic disease, the species was already persecuted for decades beforehand and survived for at least another 16 years and 9 months. Thus Paddle needs an even more devastating tragedy to have visited the mainland than Tasmania. Whose scope is barely conceivable, if the systematic killing of thylacines combined with disease did not kill off the Tasmanian population as quickly as the mainland thylacine died out. Simply put, there just isn't any such possible extinction cause that has any documentary evidence for it.

Dingo-thylacine interactions: stiff competition, continental breakfast, or no bark, no bite?⁷

While the thylacine almost certainly didn't survive on the mainland until European colonisation, it undoubtedly existed there until at least several thousand years ago. Irrefutable evidence of this are the many securely dated fossils (White et al., 2018b). The question naturally arises, why did it persist in Tasmania so much longer? Three basic hypotheses exist⁸: climate, competition with the dingo, and human intensification (population size and technological advancements). Evidence suggests that New Guinea and Tasmania were less affected by global climatic changes taking place at the time we are talking. While Tasmania almost certainly lacked canids, and seems to have lacked the spike in technological evolution that took place on the mainland. Therefore, it is very difficult, if not impossible, to single out any one of these three hypotheses as the reason they survived in Tasmanian much longer. Moreover, geologically recent extinctions can be quite complex affairs. The Tasmanian thylacine population is a case in point. No sole extinction cause can be cited, even if one of them would have been sufficient had the others not arisen.

The dingo has long been held as a cause of the thylacine's extinction on the mainland, due to what we might ambiguously call the 'superiority hypothesis'. As already noted the larger brains, bigger social structure, and broader diet of the dingo, potentially have serious implications for any sympatry between the species. Aboriginal author Marie Munkara, however, suggests that they were in general not sympatric at all⁹. Contrary to Paddle, accepting the dingo as superior to the thylacine in the relevant sense only commits us to the view that thylacines would have been more negatively affected by sympatric dingoes than dingoes would have been by sympatric thylacines. Indeed Paddle seems unfazed at calling some dogs "stupid" (Paddle, 2000:21). While it is certainly true that a strong correlation between brain size and intelligence is somewhat wanting, the implication of his comment that brain size is rather irrelevant to intelligence is even less attested.

Ogilby (1841) was the first to suggest explicit competition between thylacines and dingoes, resulting in the loss of the thylacine from the mainland. A view contradicted by Gerard Krefft's claim that the thylacine was actually responsible for the extirpation of the dingo from Tasmania, which as (Allport, 1868) points out, is rather a bold claim since the dingo is not know to have ever occupied the Apple Isle. In his PhD thesis, Robert Paddle compiled a list of references from 1842-2000 which claimed that dingoes were responsible for the extinction of the thylacine on mainland Australia (Paddle, 1997). From 1842-1950 he found 19 references, while during the next 50 years he found 80. An increase from one every 5.5 years to one every 8 months (see Paddle, 2000:35f). While one needs to factor in such things as the increase in the rate at which literature is being written and published as the population of scientists grows, this is nevertheless an abrupt transition in the general thinking.

Paddle attributes it to the growing scientific realisation that the thylacine is probably extinct, which thus calls out for a cause of its extinction. Scientists being allergic to the notion of anthropic extinction, "desperate to attribute the thylacine's extinction to non-human causes", the superiority of the dingo was constructed (whether consciously or unconsciously) as a red herring to divert attention away from the fact that we only have ourselves to blame (Paddle, 2000:20). Well let's say that Paddle is right. What logically follows from this? Certainly not the impossibility, or even improbability, of the dingo's central role in the extinction of mainland thylacines. For that you need to have contrary evidence (which Paddle claims that he has). To suggest otherwise is to fall for the genetic fallacy. That is, ruling out a view based upon not its intrinsic merits, but because of the motivations and/or reasons ascribed to those for holding it in the first place. For example, if I believed the sun revolved around the Earth simply because that is what my father told me, that would not in itself disprove geocentrism. For that you would need evidence that geocentrism itself is false. At its most basic, conclusions are independent of any particular route to getting there: disproving a route does not disprove the destination.

Paddle himself doesn't fall for the genetic fallacy. He makes it quite clear that his problem with the notion of the superiority of the dingo is, at least in part, the dissonance involved in asserting the dingo's central role in the extinction of thylacines on the mainland, while simultaneously ignoring or trying to explain away their lack of such efficacy in Tasmania. As I have already discussed, this is itself at odds with the historical record as illustrated by (Boyce, 2006,2008). Indeed as noted, Paddle's observation that the destruction and displacement of aboriginal people led to a rise in the feral dingo population on the mainland, provides a cause for the emergence of the packs of feral dogs in Tasmania that proved such a problem for thylacines during the early-to-mid 19th century. The latter only getting a reprieve after canine distemper arose. At the same time, Paddle puts forward a good point. Even if we allow that dingoes are in general superior to thylacines, were there enough of them prehistorically to actually cause the extinction of the mainland thylacine? It seems like the literature has simply assumed that there were.

Only one species of the genus Thylacinus is known to have survived into the Late Pleistocene (126ka-11.7ka) anywhere on the planet. Although in the nineteenth century two or more species were often accepted for periods before one was synonymised with the living T. cynocephalus. Either a fossil species from the recent past (e.g. T. spelaeus), or two contemporaneous species in Tasmanian (e.g. T. breviceps). On the mainland fossil female thylacines are on average somewhat smaller than their Tasmanian counterparts, a fact long known (Ride, 1964) and which prompted (Lowry, 1972) to investigate the taxonomic status of the two populations. She found no justification for the recognition of two distinct taxa. Fillios et al. (2012) further argued for the small stature of female mainland thylacines, in conjunction with a discussion of the possible implications of this for thylacine-dingo competition. While also pointing to the larger brains of dingoes, concluding that dingoes "were likely a primary agent for the extinction of the thylacine from mainland Australia".

Letnic et al. (2012) was more specific, concentrating solely upon the potential for direct killing of relatively smaller female thylacines by larger dingoes. With a skewed sex ratio, reproductive oputput would have been severely constrained, thus potentially leading to other negative effects like inbreeding. This exploits a hypothetical difference between the trophic levels of male and female mainland thylacine when the dingo is potentially sympatric, due to their differential sizes as discussed above. That is, the level at which they sit with respect to the rest of the food chain. Males would be apex predators along with male and female dingoes (since their sexual dimorphism is less marked), while female thylacines would not be apex predators. When a new, larger predator moves into an area a trophic cascade is said to occur, whereby any change at the top of the food chain affects the lower levels. In this case the smaller native predator/s all slide down the trophic scale to a lower level. Here in Australia the red fox (Vulpes vuulpes) and feral cat (Felis catus) are said to be mesopredators, that is, predators which are mid-level on the trophic scale.

Female predators tend to move less within their smaller ranges than males of the same species. Thus even if females are smaller, they are also less mobile and hence less likely to encounter species at a higher trophic level. This fact seems not to have been covered much within the literature, at least in part because it does not seem to give the thylacine much of a reprieve if dingoes are generally superior. While if dingoes are not superior, then it is a moot point. A study lead by Marie Attard (Attard et al., 2011) argued that the stress placed upon the thylacine skull during biomechanical reconstructions pointed to the thylacine probably being capable only of killing prey smaller than itself. Due largely to its relatively narrow skull, compared with a Tasmanian devil and even a quoll (Dasyurus sp.). If true this has serious implications for even a male thylacine's ability to fend itself against lone dingoes, let alone survive an encounter with a whole pack.

Johnson & Wroe (2003) argued that competition from the dingo, as well as human intensification driven by population growth which fueled technological advancements, together could have driven the three vertebrate species conspicuously absent from the mainland but present in Tasmania (thylacine, Tasmanian devil, native hen) extinct. The most recent study to date has found that thylacines and Tasmanian devils both became extinct on the mainland about 3,275 years ago (White et al., 2018b). Which would constitute a temporal overlap of about 1,200-1,700 years between the species. Although the data set used is biased towards southern Australian fossil sites, and thus does not take into account the potential for reufgia in northern Australia. Nevertheless, the relatively short overlap between the two species in southern Australia is important, even though the terminal dates for thylacines (and Tasmanian devils) on the mainland also coincides with peak ENSO activity.

If the thylacine were alive today, even if only in Tasmania, the current debate over the role of the dingo in its mainland extinction would undoubtedly look very different. For example, we would have a far better idea whether it is a pursuit, pounce-pursuit or ambush predator (Jones & Stoddart, 1998; Jones, 2003; Figueirido & Janis, 2011; Janis & Figueirido, 2014). The thylacine's stripes are suggestive of a mode of camouflage, which lends itself to an ambush style of predation. However, reports of thylacines doggedly pursuing wallabies at a steady trot tend to portray a more cursorial habit, which is somewhat inconsistent with its elbows (Figueirido & Janis, 2011). Its senses seem to have been very sharp, for example an anecdote of a pet thylacine being aware of visitors long before they arrived. While the species was evidently rarely encountered in the wild, a testament not to its rarity (since so many were killed) but rather to its ability to evade humans. In the end, it seems to have had a rather unique generalist morphology (Janis & Figueirido, 2014), indicating its true dominon over the beautiful Tasmanian landscape, not having to be shoehorned into some tight ecological niche with little evolutionary breathing space.

Quantifying the pre-European dingo population

Human colonisers often have a bifurcating effect on the fortunes of the native plants and animals of a region. Some decrease in numbers as their habitat is altered and destroyed, while for others the changes bring new opportunities. Little if any research has been done into how European colonisation changed the fortunes of the dingo. Robert Paddle suggests that the displacement and destruction of mainland aboriginals over the last 200 years is largely responsible for the dingo's current situation: "existing by themselves at high population density levels throughout the Australian mainland" (Paddle, 2000:22). There is no doubt that dingoes forced to turn feral because their owners were killed or had to abandon them contributed to a rise in the wild population. Just as the introduction of livestock onto the continent brought a greater potential food supply and more watering holes, especially in arid regions. But without a pre-European wild population baseline, it is very difficult to quantify or even infer the impact of the dingo on mainland thylacines.

Early European accounts of the dingo are too sparse in detail and too few in number to help chart the pre-European wild dingo population. While subfossil remains contribute only a small piece of the overall picture, namely minimal geographical extent. For real data on population extent and density we need aboriginal knowledge. Unfortunately the loss of many aboriginal peoples and languages means that what information is left that can be compiled is very incomplete. This information, spread over the many aboriginal tribes, has not been adequately documented. It therefore remains to be seen exactly how extensive the pre-European dingo population was. But given the huge size of Australia, the longevity of the species here, inter-tribal warfare which left dingoes without owners, the dingoes' adaptibility, and finally the pre-European abundance of prey, there is every reason to suppose that the wild population was significant. Although this remains largely conjectural.

This state of affairs is quite surprising since the whole dingo-thylacine debate potentially hinges on this very question. Even if dingoes are conclusively shown to be superior in every way to thylacine (quality), without enough of them (quantity) they could not have caused the species' extinction from the mainland. Or at least contributed their part if other factors were also involved, as is more than likely. It is as if scientists have simply assumed that the dingo's conservation status, in danger of extinction through hybridization with domestic dogs (Corbett, 2008), but more importantly were undoubtedly cosmopolitan in prehistoric times (Corbett, 1995b), means that they existed in the requisite density. In reality, distribution neither entails nor commonness nor rarity. A microendemic may be locally common, while a cosmopolitan species may be extremely rare.

New Guinea, Same Story?

The dingoes closest living relative is the New Guinea singing dog (NGSD). It is unsurprising that such a breed should exist since the dingo's ancestor was presumably brought from mainland Asia through New Guinea. However, unlike the dingo the status of the New Guinea singing dog in the wild is not well understood. It appears to be extremely rare, having been photographed only several times. At least once it has been claimed to have been rediscovered in the wild. Such claims are rather overstated as the New Guinea highlands and adjacent territories are not well surveyed. It is well known that there is a small captive population with limited genetic diversity, which owes its existence to Sir Edward Hallstrom, then president of the Taronga Park Zoo. Indeed famed Australian zoologist Ellis Le Geyt Troughton named the "species" after him: Canis hallstromi (Troughton, 1957).

The first discovery of thylacine remains from New Guinea was a part mandible excavated by Susan Bulmer in April 1960 (van Deusen, 1963), later found to be of early Holocene age (Sutton et al., 2009). Further remains, albeit far more ancient of almost certainly of a different species were reported by (Plane, 1976). More recent excavations at Nombe Rockshelter and other archaeological sites have yielded remains that are mid-Holocene in age, the youngest c.5,000 cal. BP (Mountain, 1991; but see Sutton et al., 2009). Like Tasmania, and unlike mainland Australia, New Guinea seems to have suffered far less from climatic changes at the time. Although famously the home of literally thousands of different languages, the human population was not cosmopolitan:

"At the time of European contact the population was concentrated along the coastal fringe with quite sparse and isolated groups in the rainforests and mountain slopes. Some areas, such as the Fojes Mountains, seemed to be totally without habitation." (Hope & Haberle, 2005:542)

Although unlike Tasmania, and like the mainland, it is evidently home to canids. Surprisingly, a 2016 study (Cairns & Wilton, 2016) found that Australian dingoes constitute two genetically distinct populations, with one being closer to the New Guinea singing dog than the other. This is made more surprising by the fact that the New Guinea singing dog is significantly smaller than either dingo population at the withers, which is reflected in their respective weight. This probably reflects an adaptation to their respective environments, hinting that New Guinea thylacines were even smaller than their mainland Australian coutnerparts, although New Guinea thylacines are so little studied that not even basic biometric data gathering appears to have taken place. The sheer paucity of the relevant data therefore means that a huge big question mark sits over the island. With simply nothing more to say at this point.

Conclusion

Like the broader debate over the extinction of Australia's Late Pleistocene megafauna, our understanding of the extinction of the thylacine on the mainland has evolved substantially over time. What was initially thought to be a relativey simple case of plain competition with a placental carnivore (what some would call "placental chauvinism"), has turned out to be a rather more complex affair. Our understanding of climatic changes which were taking place at the time of the thylacine's terminal decline has come leaps and bounds in the last decade or more. Which mirrors that taking place in the Americas (i.e. the Younger Dryas), and to a lesser extent Europe and Africa (e.g. Prado et al., 2015; Faith, 2014). As well as archeological evidence which suggests that human intensification could have been a significant factor in its demise.

Combined with new and more accurate dating techniques, which have helped overturn old extinction chronologies and tighten others, our picture of the Late Pleistocene-Holocene megafaunal extinctions are slowly coming together. The true cause of the thylacine's extinction on the mainland may never be fully known, however, the case is more than conjectural. We can point to explicit causes, dingo competition, climatic changes, human intensification, which may collectively have contributed to its demise. The debate is now over the relative importance of these with respect to each other. Some of which reappeared millennia later as Tasmania was visited by an even greater tragedy.

 

Appendix 1: A mounting myrmecophagous monotreme mystery

Museums are like glorified morgues. Thousands upon thousands of dead specimens sit lifeless in untold cabinets, cupboards and crates, most of which are in storage at any one time. After all proper preservation is key so that future generations can also enjoy the collections. Specimens of recently extinct species are especially hallowed, and understandably so since the loss of even a single one means that it literally cannot be replaced. Given all this, it is ironic then that museums are abuzz with activity; staff conduct research on the specimens and those on loan from other institutions, plan expeditions to acquire new specimens, seek funding to continue their many activities, and of course interact with the public in various ways. They are repositories of the natural world, amounting to our collective knowledge of the world around us. With so many specimens, many donated or bequeathed, it is hard to keep track of them all. Records today are invariably digitised, which means that you don't even have to leave your living room to go to the museum in a sense. Dead curators (not kept in museums!) would have relished the use of such technology.

There is of course something human in researching old handwritten records that does not tranfer to the digital dimension. Ink stains, different handwriting, indeed the evolution of handwriting, are all captured in dusty old books and ledgers recording the history of a museum from its humble (or not so humble) origins. While Charles Darwin died almost 140 years ago, it is perfectly possible to see specimens that he himself collected aboard the H.M.S. Beagle (1831-1836). Or to see the giant moa that famed Victorian era anatomist Richard Owen was photographed next to, one of the few instances when he was exceeded¹⁰. Incidentally, Darwin's own records of the finches he collected in the Galápagos Islands were not good enough to trace each form to its native island, so that he had to rely upon his shipmates and their more careful records (Sulloway, 1982). Moreover, museums do not always know precisely what they have. New species aren't always discovered in the field, they can also sit unrecognised in old museum collections for decades before serendipity intervenes (e.g. Bauer & Russel, 1986; Eldridge et al., 2018).

Not all discoveries made in old museum collections are new species of course. When Kristofer Helgen stumbled across a specimen of Zaglossus bruijni stated as coming from Western Australia's Kimberley region, he recognised its importance as a major extension to the recent distirbution of a monotreme, those bizarre egg-laying mammals. The genus as a whole, which also contains the extremely rare Z. attenboroughi (named after famed naturalist Sir David Attenborough), has been thought to have been historically confined to New Guinea. Such a discovery would not be unprecedented as several other species have been discovered to range over both New Guinea and Queensland's Cape York Peninsula in recent decades. However, with no other recent record of the species in Australia, one might think it more likely that there has simply been an accident whereupon the specimen has ended up with a label attached to it that doesn't belong.

Helgen and his co-authors, however, disagree. They use several lines of evidence to argue that in fact Zaglossus was a native of Australia into the recent past, and indeed may still exist. Their case in many ways mirrors that of Paddle, especially the cross-cultural aspect. There is no doubt that the species existed here in Western Australian in the past, as evidenced by fossils. However, rock art images are notoriously difficult to identify with living or recent species, since that presupposes an intent by the artist to document accurately what exists. Aboriginal art includes many depictions that everyone agrees do no represent real biological entities, and so it is somewhat arbitrary to point out morphologically ambiguous art and assign it taxonomically to a species. Contra (Helgen et al., 2012) there are no uncontroversial records of any Australian megafaunal rock art depictions (Bednarik, 2013; Lewis, 2017). If we accept that the rock art depicts an echidna it is far more likely to represent Tachyglossus aculeatus than Zaglossus bruijnii (see below).

More pertinent are the claimed aboriginal memories of two distinct species of echidna. In general native people are extremely good at distinguishing between species, which lends credence to their memories. Moreover their remote locations mean that they are often better placed to provide science with knowledge of animals than scientists are. Especially rare, extirpated or extinct taxa (e.g. Burbidge & Fuller, 1979; Burbidge et al., 1988; Tunbridge, 1991). However, methodology and interpretation are everything. And it seems that (Helgen et al., 2012) have fallen into the trap of desiring aboriginal corroboration a bit too much. After all, without it a label mix up is almost certain. After reading their paper, Andrew Burbidge made extensive inquiries with the Nykina-Mangala and Wunambal people on whose land the echidna was supposedly collected, and has found that both groups only recognise one kind of echidna (Burbidge, 2017). The couple of reports of Zaglossus knowledge by aboriginals cited by (Helgen et al., 2012) are thus anomalous.

The biggest problem, however, is that even if we accept for the sake of the argument that two species of echidna existed in Australia until at least 1901, it should still be here. And yet no other specimens have come to light either before or since, despite numerous surveys and expeditions (see Burbidge, 2017) and a $43k four-year grant. Burbidge (Ibid.) thus concludes that the "simplest and most plausible explanation" is that a label mix up occured sometime between the specimen leaving Perth and arriving in Tring (the location of Lord Walter Rothschild's collection). This is a far simpler explanation than that a species of echidna has largely gone unnoticed by both aboriginal people and science. Incidentally, Burbidge (Ibid.) recounts an instance of a Wunambal man, Geoffrey Mangolamara, telling he and colleague Phillip Fuller of the thylacine: "this was volunteered and not in response to a question from us – and he made it clear that it had not occurred in his country for many generations".

 

Appendix 2: An unfinished list of pre-1950 references to dingo extermination

Devaney, James. (1930). Nature Notes. The Marsupial Wolf. Daily Mercury (Queensland), Friday, 13 June, p. 5.

Ogilby, W. (1841). Notice of certain Australian quadrupeds, belonging to the order Rodentia. Transactions of the Linnean Society of London 18: 121-132.

 

Notes:

¹ Although unrelated to the present article, I find it fascinating enough to mention that at least two breeds of dog from northern USA and southern Canada, the Salish Wool dog and the Plains-Indian dog, were both bred for their woolly coats. Their wool was literally used to make blankets, and their extinction was in part caused by the introduction of sheep.

² There has also been a less covered discovery of a mummified thylacine head in Murra-El-Elevyn Cave, also on the Nullarbor Plain, that took place sometime in 1990. However, I lack further details.

³ R. J. W. Selleck has proposed that 'Cambrian' is the well known former museum director Frederick McCoy (Selleck, 2003:83).

⁴ This would seem to predate the earliest known transportation of live thylacines to the mainland from Tasmania. However, the animal needn't have been alive when it left Tasmania. Equally it might have died during the journey.

⁵ The fact that 'Dr.' Litchfield was a conman was brought to my attention by thylacine researchers Chela Tnyi (pseudonym) and Gareth Linnard, who each thoroughly researched his background.

⁶ It is certainly true that Holocene thylacines exhibited reduced genetic diversity (Menzies et al., 2012; White et al., 2018a). However we also have relatively little genetic diversity. Moreover, mutations would have accumulated over thousands of years, thus injecting new alleles into the mix.

⁷ For a review, see (Whelan, 2007).

⁸ Another possibility, too brief to discuss in any real detail, is that the arrival of the dingo introduced a parasite to the mainland which wiped out the thylacine (Freeland, 1993).

9 See her comments in (Judd & Steer, 2018).

10 He also came off second best against Gideon Mantell, who discovered the first dinosaur: Iguanodon, during their famous dinosaur dispute. Deborah Cadbury's book The Dinosaur Hunters (2000) gives an excellent extended account of the saga.

 

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Most Recent Global Extinctions

The following table is tentative. There are doubtless many species unknowingly omitted from consideration simply because they went extinct without ever having been documented. While other (sub)species not included here probably survived longer than is recognised by science, and possibly still exist. Moreover, the "Last record" given for each (sub)species almost certainly does not represent the very last individual (endling) in almost all cases for obvious reasons. Nevertheless such an incomplete picture of recent extinctions is better than none at all. The list is arbitrarily limited to extinctions that are believed to have taken place since 1 January 2000^1,2.

Entries listed in bold are tentative post-1999 records.

Entries which are underlined may not be taxonomically valid.

Entries which are both bolded and underlined are taxa which are feared extinct again.

An asterisk '*' denotes that the endling died in captivity/ex situ.

 

Scientific name	Common name (type of organism)	Geographical range	Last record	References
Achatinella apexfulva*	(land snail)	Oahu, Hawaiian Islands, USA	1 January 2019	David Sischo
Ecnomiohyla rabborum*	Rabb’s fringe-limbed treefrog	Panama	26 September 2016	Platt, 2016
Partula faba faba*	Captain Cook's bean snail	Raiatea, Society Islands	21 February 2016	Justin Gerlach's website
Emoia nativitatis*	Christmas Island forest skink	Christmas Island, Indian Ocean	31 May 2014	Woinarski et al., 2017
Megupsilon aporus*	Catarina pupfish	México	2014	González et al., 2018
Chelonoidis abingdonii*	Abingdon Island tortoise	Abingdon Island (=Pinta Island), Galápagos	24 June 2012	Jones, 2013
Diceros bicornis longipes	Western black rhino[ceros]	West Africa	2012	Gippoliti et al., 2018
Philydor novaesi	Alagoas foliage-gleaner (bird)	Alagoas State, Brazil	September 2011	Lees et al., 2014; Pereira et al., 2014
Brachylagus idahoensis ssp. nov.*	Columbia Basin pygmy rabbit	Columbia Basin, Washington, USA	2011 or after	Becker & DeMay, 2016
Garra festai	Ammiq garra (fish)	Ammiq Marshes, Lebanon	2011	Freyhof et al., 2014
Rhinoceros sondaicus annamiticus	Vietnamese Javan rhino[ceros]	Cambodia, Laos, Thailand and Vietnam	29 April 2010	Brook et al., 2011
Alcelaphus buselaphus tora	Tora's hartebeest (mammal)	Nubia and W Ethiopia	2010	Gippoliti et al., 2018
Pipistrellus murrayi	Christmas Island pipistrelle (bat)	Christmas Island, Indian Ocean	27 August 2009	Lumsden, 2009
Melomys rubicola	Bramble Cay melomys (rodent)	Bramble Cay, Torres Strait	2009	Woinarski & Burbidge, 2016
Tryonia oasiensis	(springsnail)	Caroline Spring, Texas, USA	2009	Hershler et al., 2011
Festuca molokaiensis	(grass)	Molokai, Hawaiian Islands, USA	2009	Wood et al., 2019
Atelopus pinangoi	Green and red harlequin toad	Venezuela	December 2008	Pereira-Muñoz et al., 2015
Asterropteryx gubbina	(goby fish)	Silhouette island, Seychelles	April 2008	Gerlach, 2010
Pseudophoxinus syriacus	Barada Spring minnow	Syria	2008	Freyhof et al., 2014
Claravis geoffroyi	Purple-winged ground dove	Argentina, Brazil & Paraguay	November 2007	Areta et al., 2009
Cichlocolaptes mazarbarnetti	Cryptic treehunter (bird)	Alagoas State, Brazil	2007	Pereira et al., 2014
Telmatobius mendelsoni	(amphibian)	Peru	2007	De la Riva et al., 2012
Plectostoma charasense	(snail)	Pahang state, Malaysia	2007	Thorseng, 2014
Bombus franklini	Franklin's bumble bee	California & Oregon, USA	9 August 2006	Kevan, 2008
Chirostoma bartoni	Alberca silverside (fish)	Alberca Caldera, Guanajuato, México	August 2006	Jelks et al., 2008
Samoana attenuata	Cone-shaped tree-snail	Society Islands, French Polynesia	February 2006	Coote, 2009a
Eleutherodactylus brevirostris	Shortsnout robber frog	Hispaniola	2006	Hedges & Díaz, 2009
Holcaspis brevicula	Eyrewell ground beetle	Canterbury Plains, South Island, New Zealand	9 February 2005	Brockerhoff et al., 2005
Podarcis filfolensis kieselbachi	Selmunett lizard	Selmunett Island, Malta	2005	Sciberras & Schembri, 2008
Nymphargus truebae	Trueb's Cochran frog	Peru	2005	IUCN ASG, 2017c
Dypsis brittiana	(palm tree)	Madagascar	2005	Rakotoarinivo & Dransfield, 2012
Melamprosops phaeosoma*	Po'ouli, Black-faced honeycreeper	Maui, Hawaiian Islands	28 November 2004	Song, 2004; Powell, 2008
Senecio berteroi	(flowering plant)	Robinson Crusoe Island, Juan Fernández Islands, Chile	August 2004	Danton and Perrier, 2005
Toxostoma guttatum	Cozumel thrasher (bird)	Cozumel, México	June 2004	Curry et al., 2006
Delamarephorura tami	(non-insect arthropod)	Viet Nam	2004	Deharveng & Bedos, 2016a
Patiriella littoralis	Derwent River seastar	Tasmania, Australia	2004	Department of the Environment, 2020
Psephurus gladius*	Chinese paddlefish	China	2004	source
Mastigodiaptomus galapagoensis	El Junco Lake copepod	El Junco Lake, San Cristóbal Island, Galápagos Islands	2004	López et al., 2021; Elías-Gutiérrez et al., 2023
Nesiota elliptica*	St Helena olive	St Helena, Atlantic Ocean	December 2003	Lambdon & Ellick, 2016
Xenopoecilus poptae	Popta's buntingi	Lake Poso, Sulawesi, Indonesia	March 2003	Parenti & Soeroto, 2003
Nectophrynoides poyntoni	(amphibian)	Udzungwa Mountains, Tanzania	Early 2003	Menegon et al., 2004
Atelopus farci	Forest stubfoot toad	Colombia	2003	IUCN ASG, 2017a
Atelopus simulatus	Imitating stubfoot toad	Colombia	2003	IUCN ASG 2015
Altiphrynoides osgoodi	Osgood's Ethiopian toad	Ethiopia	2003	IUCN ASG, 2014
Kubaryia pilikia	(land snail)	Peleliu, Palau	2003	Rundell, 2012
Bonatea lamprophylla	(orchid)	Greater St. Lucia Wetland Park, St. Lucia	2003	Combrink & Kyle, 2006
Lipotes vexillifer*	Baiji, Yangtze River dolphin	Yangtze River, China	2002 (2007?)	Turvey, 2008
Partula labrusca*	Vine tree-snail	Raiatea, Society Islands	2002	Coote, 2009b
Cereus estevesii	(cactus)	Minas Gerais State, Brazil	2002	Braun & Taylor, 2013
Glaucidium mooreorum	Pernambuco pygmy owl	Pernambuco State, Brazil	November 2001	BirdLife, 2016
Libythea collenettei	Marquesan snout butterfly	Marquesas Islands, French Polynesia	28 August 2001	Kawahara, 2003; Kawahara & Toussaint, 2018
Numenius tenuirostris	Slender-billed curlew (bird)	Northern Hemisphere	15 April 2001	Oláh & Pigniczki, 2010
Xerobdella lecomtei	European land leech	near Graz, Austria	2001-2005	Kutschera et al., 2007
Atelopus sernai	(amphibian)	Antioquia Department, Colombia	2001	IUCN ASG, 2017b
Labeobarbus reinii	Giant Atlas barbel	Morocco	2001	Freyhof & Ford, 2022
Hyperolius ruvuensis	(amphibian)	Tanzania	2001	Barratt et al., 2017
Argynnis hyperbius inconstans	Australian fritillary (butterfly)	NSW & Queensl., Australia	2001	Lambkin, 2017
Phyllostegia knudsenii	(flowering plant)	Kaua'i, Hawaiian Islands	2001	Clark, 2015; Wood, 2015; Wood et al., 2019
Tetramolopium capillare	pamakani (plant)	Maui, Hawaiian Islands	2001 (Federal Register); 2006 (Wood et al.)	Federal Register 81(61); Wood et al., 2019
Glyceria drummondii	Nangetty grass	Western Australia, Australia	2000-2004	Threatened Species Week entry
Hesperia meskei pinocayo	Rockland grass skipper butterfly	Florida, USA	2000	Mongabay
Eutamias atristriatus	Penasco least chipmunk	New Mexico, USA	2000	Suckling et al., 2004
Conturbatia crenata	(land snail)	Fregate island, Seychelles	2000	Gerlach, 2009
Samoana minuta	(land snail)	Fatu Hiva, Marquesas Islands	2000	Gerlach, 2016
Pyrgulopsis torrida	(land snail)	Ventura County, California, USA	2000	Hershler et al., 2016
Rhizopsammia wellingtoni	Wellington's solitary coral	Galápagos Islands	2000	Hickman et al., 2007
Solanum ruvu	(flowering plant)	Ruvu Coastal Forest, Morogoro District, Tanzania	2000	Vanishing Flora forum
Euphorbia celastroides tomentella	(flowering plant)	Oahu, Hawaiian Islands, USA	2000	Wood et al., 2019
Acaena exigua	(flowering plant)	Kaui & Maui, Hawaiian Islands, USA	2000	Wood et al., 2019
Capra pyrenaica pyrenaica	Bucardo, Pyrenean Ibex	Pyrénées Mountains, Europe	6 January 20003	Folch et al., 2009
Peperomia subpetiolata	(flowering plant)	Maui, Hawaiian Islands, USA	c.2000	Wood et al., 2019
Zosterops albogularis	White-chested white-eye (passerine bird)	Norfolk Island	2000	Butchart et al., 2006

Notes:

¹ The epiphytic fern Adenophorus periens was formerly listed above, last recorded in 2014 from the Hawaiian Islands. However, it was rediscovered in 2021 (source: Rare Plant Program Highlights 2021).

² The Cyprus grass snake (Natrix natrix cypriaca) was formerly listed above with a last record of >2007 (Baier & Wiedl, 2010) but has sinc been found again (Zotos et al., 2021).

³ A cloned specimen was born, and died after several minutes on 30 July 2003 (source).

 

 

Appendix 1: Collected pre-2000, but habitat destroyed post-2000 and thus probably causing extinction

Scientific name	Common name	Distribution	Last record	Habitat destroyed
Lucihormetica luckae1	Lucka's cockroach	Tungurahua volcanoe, Tungurahua Province, Ecuador	1939	2010

¹ Vršanský, P., Chorvát, D. A., Fritzsche, I., Hain, M. and Ševčík, R. (2012). Light-mimicking cockroaches indicate Tertiary origin of recent terrestrial luminescence. Naturwissenschaften 99(9): 739-749. [Abstract]

 

Appendix 2: May be extinct, but confirmation is lacking

Scientific name	Common name/type	Distribution	Last record	References
Pseudococcus markharveyi	Banksia montana mealybug	Stirling Range National Park (Bluff Knoll and Pyungorup Peak), Western Australia, Australia	November 2019	Moir, 2021
Drosera allantostigma	Geraldton sundew	Geraldton Sandplains, Western Australia, Australia	December 2018	Cross, 2022
Saxicolella deniseae	(plant)	Guinea	27 January 2018	Cheek et al., 2022a
Sitta insularis	Bahama nuthatch	Grand Bahama, the Bahamas	2018
Microcambeva bendego	(catfish)	Rio de Janeiro, Brazil	31 August 2016
Inversodicraea senei	(plant)	Memve’ele Falls, Ntem River, Cameroon	2016	Budden & Cheek, 2022
Spathoglottis arunachalensis	(plant)	Arunachal Pradesh, India	2016	Tsering, 2022
Mollinedia myriantha	(plant)	Rio de Janeiro, Brazil	2015	de Lirio et al., 2022 [preprint]
Aegiphila caymanensis	(plant)	Grand Cayman, Cayman Islands	2015
Musa paramjitiana	(wild banana)	Andaman & Nicobar Islands, India	2013
Sulawesidrobia yunusi	(snail)	Sulawesi, Indonesia	2013	Rintelen, 2019a
Tylomelania zeamais	(snail)	Sulawesi, Indonesia	2013	Rintelen, 2019b
Ischnura solitaria	(dragonfly)	Colombia	2013	Bota-Sierra & Sandoval-H, 2021a
Eremospatha barendii	Rattan, Rattan palm	Cameroon	2012	Cosiaux et al., 2016
Curcuma pygmaea	(plant)	Vietnam	17 July 2010	Leong-Škorničková & Tran, 2019
Tsoukatosia evauemgei	(snail)	Peloponnesos, Greece	July 2010	Reischütz, 2017
Epigomphus wagneri	(dragonfly)	Costa Rica	27 June 2010	Bota-Sierra et al., 2021a
Epigomphus morrisoni	(dragonfly)	Costa Rica	22 June 2010	Bota-Sierra et al., 2021b
Craugastor fleischmanni	Fleischmann's robber frog	Costa Rica	2 March 2010
Maoridrilus felix felix	(earthworm)	South Island, New Zealand	2010
Maoridrilus felix vallis	(earthworm)	South Island, New Zealand	2010
Phrynobatrachus njiomock	Lake Oku puddle frog	Mount Oku, Cameroon	2010
Hypsolebias splendissimus	(fish)	Bahia, Brazil	2010	Lyons, 2021a
Manihot stellata	(plant)	Bolivia	2009	Fabriani, 2021
Barbodes lindog	(fish)	Lake Lanao, Lanao Province, Mindanao Island, Philippines	2008
Telmatobius bolivianus	(frog)	Bolivia	2007	IUCN SSC Amphibian Specialist Group, 2020a
Barbodes sirang	(fish)	Lake Lanao, Lanao Province, Mindanao Island, Philippines	2007
Artemisia kauaiensis	(plant)	Kauaʻi, Hawaiian Islands, USA	2006	Clark, 2016
Ceratophysella sp. nov. 'HC'	(non-insect arthropod)	Viet Nam	2006	Deharveng & Bedos, 2016b
Heteragrion archon	(damselfly)	Venezuela	2006	Vivas Santeliz et al., 2021
Musa yamiensis	(banana)	Taiwan	2006	Allen & Plummer, 2020
Senna suarezensis	(plant)	Madagascar	2006	Ramanantsialonina, 2019
Sulawesidrobia datar	(snail)	Sulawesi, Indonesia	2005	Rintelen, 2019c
Cynolebias elegans	(fish)	Bahia, Brazil	2005	Lyons, 2021b
Puccinellia gussonei	(grass)	Sicily, Italy	2005	Troìa & Peraza Zurita, 2018
Nannophryne cophotis	Paramo toad	Peru	2005	IUCN SSC Amphibian Specialist Group, 2018
Chondrostoma orientale	Oriental nase, Kor nase (fish)	Kor River basin, Iran	2005
Andinobates viridis	Green poison frog	Colombia	2005	IUCN SSC Amphibian Specialist Group, 2017d
Monanthotaxis bali	(plant)	Bali Ngemba, Cameroon	11 April 2004	Cheek et al., 2022b
Cyclophyllum tiebaghiense	(plant)	New Caledonia	2004	Barrabé et al., 2019
Orionothemis felixorioni	(dragonfly)	Bahia, Brazil	2004	Bota-Sierra & Sandoval-H, 2021b
Telmatobius sibiricus	(frog)	Bolivia	2003	IUCN SSC Amphibian Specialist Group, 2020b
Tristaniopsis jaffrei	(plant)	New Caledonia	2002	Amice et al., 2020
Psychotria torrenticola	(plant)	Cameroon	2002	Lovell & Cheek, 2021
Notropis marhabatiensis	(fish)	Mexico	2002	Domínguez, 2019
Allobates mcdiarmidi	(frog)	Bolivia	2001-2005	IUCN SSC Amphibian Specialist Group, 2020c
Kihansia lovettii	(plant)	Udzungwa Mountains, Tanzania	28 April 2001	Cheek, 2004 "2003"
Kupea jonii	(plant)	Udzungwa Mountains, Tanzania	28 April 2001	Cheek, 2004 "2003"
Allobates ranoides	Llanos rocket frog	Colombia	2001	IUCN SSC Amphibian Specialist Group, 2020d
Ochna braunii	(plant)	Tanzania	2001	Gereau et al., 2020
Sorocephalus crassifolius	Flowerless clusterhead	Western Cape Province, South Africa	2001	Rebelo et al., 2020
Oxalis hygrophila	(clover plant)	Western Cape Province, South Africa	2001	Dreyer et al., 2012
Atelopus angelito	Toad	Colombia	2000
Ophthalmolebias perpendicularis	(fish)	Bahia, Brazil	2000	ICMBio, 2022
Speonectes tiomanensis	(fish)	Gua Tankok Air cave, Gunung Kajang, Pulau Tioman, Peninsular Malaysia	2000	Ahmad, 2020
Taudactylus rheophilus	Northern tinker frog	Queensland, Australia	2000	IUCN SSC ASG, 2022
Gentiana wingecarribiensis	Wingecarribee gentian, Swamp gentian	New South Wales, Australia	2000	Silcock et al., 2021
Wikstroemia skottsbergiana	Skottsberg's wikstroemia, Skottsberg's false ohelo, ‘Ākia	Hanalei Valley & Kauhao Valley, Kauai, Hawaiian Islands, USA	January 2000	Wolkis, 2018

 

Appendix 3: Most likely extinctions in the near future^1,2

Scientific name	Common name/type	Geographical range	Number of individuals	References
Merulaxis stresemanni	Stresemann’s bristlefront	Bahia, Brazil	1 female	source
Bubalus quarlesi	Mountain anoa	Sulawesi, Indonesia	1 male	source
Hyophorbe amaricaulis	Loneliest palm	Mauritius	1 plant	source
Ceratotherium simum cottoni	Northern white rhino	East and Central Africa	2 females	source
Rafetus swinhoei	Yangtze giant softshell turtle	China	2 males, 1 sex unknown	source
Phocoena sinus	Vaquita porpoise	Gulf of California, Mexico	Unknown	source

¹ The Sehuencas water frog (Telmatobius yuracare) was formerly listed here, as there was only a single individual known ('Romeo'). The species had not been seen in the wild for 10 years or more. In late 2018 or early 2019 five further specimens were located in the wild.

² The Rufous-fronted laughingthrush was formerly listed here, but it has since been rediscovered in the wild.

 

Appendix 4: Extinct in the wild since 2000 [in progress]

* = tentative

Scientific name	Common name/type	Distribution	Last record
Telmatobius dankoi*	Loa water frog	El Loa province, Chile	June 2019
Schiedea haakoaensis	(plant)	near Laupāhoehoe, Hawaiʻi (=Big Island), Hawaiian Islands, USA	2019
Phyllostegia pilosa	(plant)	Maui & Molokai, Hawaiian Islands, USA	2018
Phyllostegia helleri	(plant)	Kauai, Hawaiian Islands, USA	2018
Cyanea truncata	(plant)	Oahu, Hawaiian Islands, USA	2018
Gracula religiosa miotera	Simeulue Hill myna (bird)	Sumatra, Indonesia	2017-2018
Phyllostegia bracteata	(plant)	Maui, Hawaiian Islands, USA	2017
Phyllostegia mannii	(plant)	Molokai & Maui, Hawaiian Islands, USA	2016
Phyllostegia brevidens	(plant)	Hawai'i & Maui, Hawaiian Islands, USA	2016
Deparia kaalaana	(plant)	Hawai'i, Kauai & Maui, Hawaiian Islands, USA	2016
Delissea rhytidosperma	(plant)	Kauai, Hawaiian Islands, USA	2014
Dendroseris gigantea	(plant)	Alejandro Selkirk Island, Juan Fernández Archipelago, Chile	March 2014
Stenogyne kanehoana	(plant)	Oahu, Hawaiian Islands, USA	2013
Stenogyne bifida	(plant)	Molokai, Hawaiian Islands, USA	2013
Kanaloa kahoolawensis	(plant)	Kaho'olawe, Hawaiian Islands, USA	2013
Caridina dennerli	(shrimp)	Sulawesi, Indonesia	2013
Phyllostegia parviflora glabriuscula	(plant)	Hawai'i, Hawaiian Islands, USA	2012
Lachanodes arborea	(plant)	St Helena	2012
Commersonia erythrogyna	Trigwell's rulingia (plant)	south-western Western Australia, Australia	2012
Copsychus stricklandii barbouri	(passerine bird)	Maratua island, Borneo	2011
Phyllostegia parviflora lydgatei	(plant)	Oahu, Hawaiian Islands, USA	2010
Cyanea grimesiana grimesiana	(plant)	Moloka'i & Oahu, Hawaiian Islands, USA	2010
Schiedea jacobii	(plant)	Hawai'i, USA	2009 or 2010
Atelopus zeteki	Panamanian golden toad	Panama	late 200X's
Stenogyne campanulata	(plant)	Kauai, Hawaiian Islands, USA	2008
Phyllostegia kaalaensis	(plant)	Oahu, Hawaiian Islands, USA	2008
Cyanea grimesiana grimesiana	(plant)	Koʻolau Mountains, Oʻahu, Hawaiian Islands, USA	2008
Magnolia wolfii	(plant)	Risaralda Department, Colombia	August 2006
Musa zaifui	(banana)	Yunnan Province, China	2006
Aphanius sirhani	(fish)	Azraq Oasis, Wadi Sirhan valley, Syrian desert, Jordan	2004-2006
Aphanius saourensis	(fish)	Oued Saoura Basin, Sahara desert, Algeria	2004
Encephalartos brevifoliolatus	(cycad)	Limpopo Province, South Africa	2004
Costus vinosus	(plant)	Panama (central)	2004
Nectophrynoides asperginis	Kihansi spray toad	Kihansi Falls, Kihansi Gorge, Udzungwa Mountains, eastern Tanzania	2004
Apricaphanius saourensis	Sahara killifish	Oued (River) Saoura River, western Algeria	2003 or 2004
Cyanea superba superba	(plant)	Oahu, Hawaiian Islands, USA	2003
Cyanea pinnatifida	(plant)	Oahu, Hawaiian Islands, USA	2001
Anaxyrus baxteri	Wyoming toad	Laramie Basin, Wyoming, USA	2000's
Clermontia peleana peleana	(plant)	Hawaii (=Big Island) & Maui, Hawaiian Islands, USA	2000

Source for Hawaiian plants: Wood et al., 2019 (excluding Clermontia peleana peleana and Schiedea haakoaensis). Delissea argutidentata was formerly listed here (last wild record 2002) before its rediscovery in the wild in early March 2021.

 

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Acknowledgements

I thank Jonathan Oliver for pointing out that I had failed to remove the Sehuencas water frog (Telmatobius yuracare) from Appendix 3.

 

Defining Extinction

By Branden Holmes

Extinction is a tricky term to define. Especially since it now has usage outside of a biological context. At the level of the individual organism the term 'death' is invariably used, although 'extinction' is considered a valid synonym. Above the level of the individual, from (sub)population level upwards, the term 'extinction' is invariably used, although 'death' is considered a valid synonym (e.g. the death of the dinosaurs). It seems then that 'extinction' and 'death' are synonymous at every biological level, which begs the question of why we have two terms to describe the exact same range of phenomena.

Other such semantically equivalent pairs of terms can be found in the English language, products of the vagaries of linguistic evolution. The question then becomes, should we make a concerted effort to distinguish them semantically, or allow them to retain their common meaning? Obviously both options have their benefits and their drawbacks, and I as an individual have no vested authority to make the decision myself. However, as nobody is under any obligation to accept anything that I write, I feel free to make a suggestion, which readers (and others) may either choose to accept, reject or ignore.

I submit the suggestion to limit the term death to the level of the individual, and to reserve the term extinction for those biological levels above the individual. The logic behind this submission is that while death and extinction are both natural occurences, the former has a much stricter timeline. Admittedly death and extinction (as I have just defined them) can both occur 'prematurely', which itself needs carefully defining, but the former is restricted to a much tighter chronology. After all, although it varies by several orders of magnitude from species to species, it remains relatively stable amongst the individuals of a single species. And even if the odd individual survives much longer than its conspecifics, its lifespan is not indefinite.

Whereas for (sub)populations there is no defined upper limit to their longevity since they are not tied to any particular taxon, and thus cannot become phyletically extinct (i.e. extinction via intra(sub)population genotypic/phenotypic replacement). Whereas for species and subspecies, although there is technically no defined temporal upper limit to their longevity, there is such a limit genotypically/phenotypically. Although a largely arbitrary delineation, (sub)species can only evolve so much before they cease to remain the same taxon. Thus in one sense they can become extinct simply through evolution, whereby although they have living descendents, all of the members of the population/s no longer express the diagnostic traits that the taxon was described on the basis of (i.e. phyletic extinction). Exactly how many diagnostic traits the (sub)population/s can or must lose before the evolutionarily immediately prior taxon is considered extinct is taxonomically subjective.

Published 7 August 2017.

 

 

Conservation Status is Onto-Epistemic

By Branden Holmes

The IUCN RedList is one of the world's most cited conservation-orientated resources. But as powerful a tool as it is, it also, like almost anything, has its shortcomings. Often assessments are not updated in a timely matter, taxa are not (yet) assessed at all, and then taxa which were rediscovered years ago are sometimes still listed as 'Extinct'. But even if the RedList were perfect, even if all available information were compiled and used as the basis of all conservation assessments, it would still not be perfect. That is because conservation status is onto-epistemic, not ontic.

This website is an attempt to compile all available information on every single taxon that meets any two (or more) of a small number of criteria:

Necessarily must meet this criterion:

• Believed to have survived until the start of the Late Pleistocene (c.126ka)

Meets at least one of the following additional criteria:

• Believed to be extinct, probably extinct, or is "missing" (50% chance it is extinct, or greater)
• Believed to have been rediscovered, either globally or in the wild, after meeting the above criterion
• Currently believed to be extinct in the wild
• Formerly considered to be a valid taxon/breed/variety that meet at least one of the above criteria

Notice that every single criterion mentioned is actually predicated upon human beliefs and knowledge (i.e. epistemology), rather than actual ontic status. That is to say, the conservation status assessment of any given taxon is derived not from any automatically corrective correspondence with reality (to mention only one of many theories of truth). Rather it is from data gathered by the world's humans. Some or all of that data may be inaccurate, misleading or just plain wrong. We humans are subjective beings, the fallible genetic and experiential products of a truly digital world (i.e. the biological world, where DNA is essentially digital code) in which the inexorable increase of entropy (as a rule of thumb) in the wider physical universe has been temporarily reversed provincially¹.

The Thylacine: A Case Study

At the start of the Holocene epoch 11.7ka, the thylacine is known to have existed in what is modern day New Guinea, mainland Australia, Tasmania as well as some other small Australian islands (i.e. Kangaroo Island, South Australia and Hunter Island, Tasmania). We know this because their palaeontological and recent remains have been found there. But just because we have not found their remains in other places does not mean that they were necessarily limited geographically only to these areas. Nor is there a guarantee that the youngest remains that we have for the species in each of these areas represents the very last individual/s inhabiting those areas.

In Tasmania in particular, since the species survived there so recently (until at least 7 September, 1936), and assuming that it is now sadly extinct there, the most recent Tasmanian remains are far more likely to be of some of the very last individuals than, say, those from New Guinea known from the mid-Holocene (c.5-5.5ka). Now of course, barring what would be the truly remarkable discovery of an entirely new population of living thylacines, I previously made the assumption, as part of my argument, that the thylacine is now extinct in Tasmania. Of course this is not necessarily the case.

The general scientific opinion is that the species is indeed wholly extinct throughout its former range. However, that statement is qualified for good reason. Not even the most ardent critic of claims of an extant population are claiming that it is impossible for the species to still exist. Rather, the thylacine almost certainly (or probably, or likely etc.) does not exist. There can be no reasonable doubt that the thylacine once existed, whatever its current fate. Thus it cannot be the case that thylacines cannot survive because it is impossible for them to do so on the basis that they are biologically impossible. Nor even that insufficient habitat remains for them. One trip to Tasmania (or just about anywhere in New Guinea, and indeed many places on the mainland of Australia) will quickly dispel that notion.

Therefore, when assessing the conservation status of the species, the aim is to try and model the most accurate representation of the species' actual ontic status in the wild and/or captivity. This involves things like the most recent record of the species, estimates of population sizes at various times, the methodological appropriateness of expeditions/surveys, etc. Notice that it is an implicit assumption in all of this that the species could still exist from a metaphysical point of view. What the compilation of available data and hermeneutical tools do for us is allow us to assign the species to one of any number of different conservation categories, depending upon what the final analysis turns out to be.

The Conservation Categories Themselves

The IUCN RedList utilises a range of conservation categories such as 'Vulnerable', 'Endangered' and 'Critically Endangered'. Most of these categories are what we might call onto-epistemic categories. They are a hybrid between ontology and epistemology. They represent our beliefs about the ontic status of the assessed taxon. That is, they attempt to say something about us but also something about external reality (in this case, reality beyond the conjunction of all humans²). They speak to our beliefs/knowledge of the assessed taxa. Of course there is one exception. The category 'Data Deficient'. This is a purely epistemic category, one that speaks to the paucity of data, and therefore our ignorance, and therefore our inability to confidently assign the taxon to any particular category that says something about the taxon itself and the health of its global population. It is a position of agnosticism.

Conclusion

When one speaks of the conservation status of a (sub)population, (sub)species etc. one can claim knowledge of a taxon's true ontic status. Or one can simply mean that given the balance of evidence, the most likely status of the wild/captive global population is this or that category. In general what is meant is the latter. After all, the former strays as much into philosophy as it does biology. The issue of whether true knowledge (i.e. cartesian certainty) is even metaphysically possible is highly contentious. One can of course retort that such a definition of knowledge is far too strict, and that the term is more often used to mean something epistemically much less strict. But then this kind of 'knowledge' starts look awfully like the evidential position espoused by the latter approach.

Notes

¹ One of the definitions of 'provincial' is a '[general] lack of culture'. The provincial reversal of entropy (i.e. the origin and evolution of life) means that that lack of culture is no longer the case. Most people would consider bacteria as primordial life, and a culture (i.e. special kind of group) of bacteria being plural instead of the singular represents the increase of life. Bacterial culture, now routine research tools in medicine (which combines both epistemology and ontology (the subject of this article), the study of the process that created us. Like the illustration of the eye sat atop the left-hand column of the 'U' shape that is able to gaze back at the process that it is the end product of. Although I wouldn't wish to affirm any speciesistic notion that we are in any way superior to any other form of life, despite mainly religious claims to the contrary). An ultra punny element for those who desire it.

² Technically, 'humans' covers any member of the genus Homo. But since we are the only extant taxon I am using the term as a synonym of Homo (sapiens) sapiens.

Published 5 August 2017.