The largest TEC ichnofossil scratch marks can only have been made by T. carnifex. Smaller marks may be attributable to a number of agents, but juvenile T. carnifex seems the most likely given the known biology of species present and comparisons with trogloxene behaviour elsewhere. The smaller TEC claw marks bear a cursory resemblance to the longer descent marks made by koalas, but the sheer number of marks is more consistent with one or more species of trogloxene rather than individuals of an obligate arboreal folivore that may have unwittingly entered the cave. While common brushtail possums are opportunistic trogloxenes, the light marks that they habitually leave on bark and on cave surfaces22 are not consistent with the deep scratches observed in TEC. Use of caves for shelter is known for rock-wallabies, thylacines and devils23,24, but while rock-wallabies are highly adept in steep terrain, they primarily use their hind limbs to move ricochetally. No TEC marks match the broad macropodid pedal claw morphology. Rock-wallabies are capable of transferring weight to their forepaws to pivot on ledges during ascent of rocky slopes, but most of the weight is borne on the pads. The large volume and broad distribution of TEC claw marks are more consistent with primarily quadrupedal animals for which manual claw use is critical. Only some aspects of the biology of the thylacine were documented prior to its extinction in 1936, but there is no record of climbing and its limbs lack any features indicative of a scansorial ability25. By contrast, devils, especially juveniles, are capable climbers23.

The modern devil IDS range coupled with the observation that Pleistocene S. harrisii, like several of its contemporaries, was smaller in the west26, suggest that only marsupial lions could have generated scratch sets with IDS values >14 mm. Devils could feasibly have contributed to the smaller end of the spectrum. However, IDS ranges for the modern species are normally distributed whereas the TEC range, although unimodal, is strongly positively skewed (Fig. 3). This could imply that the claw-mark assemblage was produced by both marsupial lions and devils, or that marsupial lions alone produced the assemblage, with most claw marks made by juveniles. The latter would be consistent with use of the cave as a maternal den to protect and raise young as observed in hyaenas and large dasyurid marsupials, e.g., devils and spotted-tailed quolls (Dasyurus maculatus)23,27,28. Similar suggestions have been made on ichnofossil scratch marks in a Romanian cave, where a range of scratches of width 5–12 cm (across 4 digits) was interpreted as indicating cave bears of different ages17.

Further insight is provided by the taphonomic analysis, which reveals minimal evidence of carnivore tooth damage. Devils either accumulated a small fraction of bones themselves or, more likely, occasionally modified the remains of animals that entered the cave via a different agency. Feeding studies of devils show that they ingest most bone at or near a kill or scavenge site; an abundance of fragments derived from scats is the hallmark of their cave dens29,30. The rarity of chewed bones and absence of scat-derived fragments reinforces the view that devils were a negligible accumulator31. The lack of T. carnifex tooth marks might also reflect their minimal role in collecting bones: the relatively complete preservation of most bones, broad size range of taxa and high species richness has been used to interpret the deposit as a pitfall accumulation, with animals falling through now-blocked solution pipes along with the sediments that entombed them16. This situation parallels that seen in a number of European caves where cave bears, hyaenas and lions often occupied caves as fossil remains accumulated17,32. The differential evidence of den use also parallels that in Australia: scavenging devils and hyaenas leave traces on chewed bones; Thylacoleo and cave bears leave scratch marks on cave surfaces; thylacines and lions are principally inferred from faunal remains17. The lack of tooth marks on TEC bones attributable to T. carnifex, however, may also reflect the morphology-based interpretation of this species as a flesh specialist7,10. Tooth marks on bones attributable to marsupial lions are only sporadically encountered and apparently incidental to dragging about carcasses or stripping them of meat and viscera7,33. This is exemplified today by lions, where incidental marking of bone during flesh stripping is common at kill sites, but rarely substantial enough to be seen in the fossil record34.

The most parsimonious interpretation of the TEC evidence is that marsupial lions were primarily responsible for the claw-mark assemblage. Devils clearly used the cave at times during the depositional interval of 140–30 kyr ago, but it is improbable that the two species cohabited the chamber given the intense interspecific antagonism displayed by carnivores, especially in cave settings17. With an estimated adult body mass of 80–100 kg, T. carnifex was 10–15 times heavier than S. harrisii (6–8 kg), which, even in a juvenile-biased mob, would likely have been at a major competitive advantage within a prime den environment. In addition, despite the number of documented devil dens and known devil climbing behaviour, no den scratch marked by devils has yet been documented. The palimpsest-like nature of the TEC trace-fossil assemblage does not allow us to tightly constrain the span over which claw marks accumulated, but it may be as great as 90 kyr, given that the latest record of T. carnifex within the TEC bone deposit is in unit E, dated to 51 ± 2 kyr16. In addition, archaeological marks on moonmilk have been shown to degrade over time35, so the fidelity of marks on this material suggests no great age for the marks.

The distribution of claw marks indicates a significant bias toward juveniles. One possibility is that, at any one time, the chamber was occupied by a lone mother looking after a single litter, as in D. maculatus, where a female may raise up to six young in a cave den28. Alternatively, the communal arrangement exemplified by brown or spotted hyaenas involving multiple reproductive females36 may be a better model. The latter draws circumstantial support from the now-largely-destroyed Komatsu Cave in southeastern Australia, which preserved a series of articulated partial skeletons, including a directly associated putative adult female and juvenile individuals and a claw-mark assemblage14,37,38.

Marsupial lions, like all marsupials, would have given birth to extremely underdeveloped young that could not be left alone until becoming at least partially weaned. Adult female body mass and time to weaning is correlated in marsupials39. For thylacines (estimated adult body mass 15–35 kg), young spent three months in the pouch followed by a further month of semi-independence24. Marsupial lion females were at least three times the adult body mass of thylacines, which suggests that they may have borne young in the pouch for a minimum of four months. Carrying older pouch young while hunting probably constrained predatory efficiency or prey size range. Even without invoking the difficulties involved in carrying pouch young, mere accompaniment by cubs has been implicated in 16% of failed chases by cheetahs (Acinonyx jubatus)40. As in numerous extant carnivores, adult marsupial lions likely left semi-independent young to shelter in the cave while they went off to hunt before returning to bring food and to rest. This is precisely what is seen in living S. harrisii: once weaned, juveniles are left in a lair while the mother hunts41. The TEC claw-mark size distribution shows that, as individuals grew, they spent less time in the cave, presumably venturing out increasingly to learn from adults and contribute to hunting.

Many claw marks within TEC are located on steep surfaces, despite more gradual inclines being available on other sides of the central rock pile and boulder (Fig. 1). This suggests regular, confident, purposeful climbing with a high degree of agility. Climbing has similarly been invoked for a small number of Ursus spelaeus scratch marks high on a Romanian cave wall17. This distribution reinforces the argument, based on skeletal morphology, that T. carnifex could climb trees7,37. This is in spite of its large size and “bear-like” build10, which have been used to argue against its arboreal adeptness9. Looking at the ecologically comparable carnivorans, small size does correlate with climbing ability, but 7 of 61 species studied weighed 80kg or more and were considered capable climbers42. The density of marks on the central rock pile also points toward the most feasible Pleistocene entrance. Above the eastern end of its apex is a now-blocked, 0.7-m-wide solution pipe, the lowermost 1 m of which is clear of sediment and slopes at approximately 45° to the ground surface 6 m above. Subsidence, possibly prior to the deposition of units H and J (40–30 ka), lowered the central rock pile31 such that its apex is now 3 m below the solution pipe, prohibiting animals from exiting the cave. Prior to this time the cave would have acted as a single large chamber, accessible to the surface by marsupial lions, while acting as a faunal trap for species unable to climb out, more similar to a European open hyaena cave than the deep hibernation chamber favoured by cave bears32.

Marked variance in scratch-set orientation on steep surfaces reflects substantial limb mobility consistent with the high degree of abduction inferred from functional morphological analysis7. Similarly, the convergence and divergence exhibited by many scratch sets points to greater lateral movement of digits II–IV than has been suggested from analysis of metacarpal–phalangeal articular morphology43. These attributes would have combined to aid in gaining traction during climbing. This again is consistent with the condition in carnivorans, where more curved claws and higher metacarpal/phalanx ratio are associated with climbing taxa, assisting in providing a broad grip42. Extensive claw marking on horizontal surfaces atop the TEC central rock pile indicates that claw engagement with the substrate was able to be maintained when not climbing. By contrast, fossil manus prints of T. carnifex from a lacustrine environment in southeastern Australia preserve four distinct impressions made by digits II–V15. The absence of claw marks there shows that claws could also be raised when moving on a relatively flat surface. Thus, claw engagement when moving across flat surfaces was likely context-dependent and perhaps more likely to occur in a more unstable or dark environment. The paucity of sets composed of >3 claw marks and in particular those where one of the outside IDS mean values is substantially greater than the others, reflects infrequent use of digit I during climbing within the cave. This observation and the fact that the southeastern cave wall manus prints lack digit I impressions (see Supplementary Fig. 7), supports the suggestion, based on manual morphology, that digit I could be raised during terrestrial locomotion43, likely coming into play more when grappling with prey or tree climbing. This feature also highlights a major difference between the scratch marks made by cave bears and marsupial lions where the former almost exclusively leave indications of four digits17, whereas the latter rarely left sets of >3 marks. This may further reflect the more dynamic manus arrangement in T. carnifex, compared to the more static cave bear paw.

The TEC trace-fossil evidence shows that T. carnifex was highly proficient at negotiating a dark, complex environment. Most claw marks are on highly heterogeneous or steep surfaces, providing clear behavioural evidence to support morphological studies inferring that T. carnifex could climb trees37,43. That claw marks of younger individuals dominate the assemblage indicates that TEC was utilised for raising and protecting young over an extended duration. This supports an idea advanced half a century ago29, but hitherto unverified, that marsupial lions used caves as dens where available, which likely explains the noted overabundance of their remains in cave skeletal assemblages11,14.

Dens are used by hyaenas to assist raising of young over extended periods of time. For hyaenas this is facilitated by their large groups, which help to defend resources from other groups44. This reflects that hyaena groups are strongly related, so that the shared costs of fights through fatalities and injuries are outweighed by the benefit to the group as a whole44. In contrast, solitary or small families of bears use caves primarily during the winter for hibernation, but also as cool places in summer, while having additional security benefits45. The element of security is one factor uniting these two ecologically distinct trogloxenes, as well as fitting palaeontological reconstructions from Europe17. Caves provide a safe, temperature controlled environment and would hence be a sought after resource. With a high proportion of juvenile marsupial lions in TEC, communal living as is seen today in hyaenas seems the most plausible arrangement to defend this resource from both other denning marsupials (thylacines and devils) as well as other conspecific groups.

The absence of tooth marks on TEC bones attributable to T. carnifex reinforces the idea the deposit represents a pitfall accumulation and is consistent with the dentition-based interpretation of T. carnifex as a flesh specialist. The high bite strength and advanced meat-slicing capabilities of T. carnifex represent the most extreme manifestation of the tendency for large, felid-like taxa to evolve where felids are absent46. By comparison, the second-largest Australian mammalian carnivore (thylacine) hunted individually or in pairs and focused on smaller prey47. Given that marsupial lions were apparently adapted to apprehending and consuming large prey10 and potentially social, it is feasible that, as in all extant group-living mammalian predators35, they were cooperative hunters. As body mass, group living and group hunting are seen as co-adaptations for procuring large prey48, it is plausible that marsupial lions were pack hunters. Such a strategy would have allowed them to prey upon the largest marsupial, the rhinoceros-sized Diprotodon optatum, bones of which have been found with incidental marsupial lion tooth marks33.