View source for Wakaleo

{{Short description|Extinct genus of marsupials}}
{{Use dmy dates|date=November 2023}}
{{Automatic taxobox
| image = Life reconstruction of the marsupial lion Wakaleo oldfieldi.jpg
| image_caption = Life reconstruction of the [[type species]], ''Wakaleo oldfieldi''
| fossil_range = {{Fossil range|Late Oligocene|Late Miocene}}
| taxon = Wakaleo
| authority = [[W. A. Clemens|Clemens]] & [[M. Plane|Plane]], 1974<ref name="Clemens1974">{{cite journal |last1=Clemens |first1=W.A. |last2=Plane |first2=M. |title=Mid-Tertiary Thylacoleonidae (Marsupialia, Mammalia) |journal=Journal of Paleontology |date=1974 |volume=48 |issue=4 |pages=653–660 |issn=0022-3360}}</ref>
| type_species = {{extinct}}'''''Wakaleo oldfieldi'''''
| type_species_authority = Clemens & Plane, 1974
| subdivision_ranks = Species
| subdivision = 
*{{extinct}}'''''W. alcootaensis''''' <small>Archer & Rich, 1984</small>
*{{extinct}}'''''W. oldfieldi''''' <small>Clemens & Plane, 1974 ([[type species|type]])</small>
*{{extinct}}'''''W. pitikantensis''''' <small>Rauscher, 1987</small>
*{{extinct}}'''''W. schouteni''''' <small>Gillespie, Archer & Hand, 2017</small>
*{{extinct}}'''''W. vanderleuri''''' <small>Clemens & Plane, 1974</small>
| synonyms = 
*''Priscileo pitikantensis'' <small>Rauscher, 1987</small>
}}

'''''Wakaleo''''' ([[Diyari language|Diyari]] ''waka'', "little", "small"; and Latin ''leo'', "lion")<ref name="Clemens1974"/> is an extinct [[genus]] of medium-sized [[Thylacoleonidae|thylacoleonids]] that lived in Australia in the Late Oligocene and [[Miocene]] Epochs.

==Taxonomy ==
''Wakaleo'' was erected in 1974 by [[W. A. Clemens]] and [[M. Plane]].<ref name="Clemens1974" /> Five species are known:

*''Wakaleo alcootaensis'' was found in the Miocene [[Waite Formation]] in the [[Northern Territory]] in 1974. It was slightly larger than the other two species.{{clarify|date=February 2020|reason=Which two species?}}
*''Wakaleo oldfieldi'' was found by a group of scientists working in the Miocene [[Wipijiri Formation]] in southern Australia in 1971. They found a nearly complete left dentary which included a few well-preserved teeth.
*''Wakaleo pitikantensis'' described by Rauscher as the type species of the genus ''Priscileo'' in 1987<ref name=":0">B. Rauscher. 1987. Priscileo pitikantensis, a new genus and species of thylacoleonid marsupial (Marsupialia: Thylacoleonidae) from the Miocene Etaduma formation, South Australia. ''Possums and Opossums: Studies in Evolution'' '''2''':423-432</ref> and placed with this genus in a revision published in 2017.<ref name="Gillespie2017">{{cite journal |last1=Gillespie |first1=A.K. |last2=Archer |first2=M. |last3=Hand |first3=S.J. |authorlink1=Anna K. Gillespie |authorlink2=Mike Archer (paleontologist) |authorlink3=S. J. Hand |title=A new Oligo–Miocene marsupial lion from Australia and revision of the family Thylacoleonidae |journal=Journal of Systematic Palaeontology |date=6 December 2017 |volume=17 |issue=1 |pages=59–89 |doi=10.1080/14772019.2017.1391885|s2cid=90758394 |url=https://figshare.com/articles/dataset/A_new_Oligo_Miocene_marsupial_lion_from_Australia_and_revision_of_the_family_Thylacoleonidae/5677702 |url-access=subscription |hdl=1959.4/unsworks_50134 |hdl-access=free }}</ref>
*''Wakaleo schouteni'' Gillespie, Archer & Hand, 2017, a mid-sized species of the genus.<ref name="Gillespie2017" />
*''Wakaleo vanderleuri'' was first found by field workers in 1967 in the Miocene [[Camfield Beds]] in the Northern Territory. Many more specimens have been found since then.

== Description ==
''Wakaleo'' is a genus of the thylacoleonid family of predatory mammals, which are known as marsupial lions. The size of ''Wakaleo'' species increases over the course of the evolution of the genus. ''W. schouteni'' is estimated to have weighed approximately {{Convert|23|kg|lb}}, comparable to a dog,<ref name="Gillespie2017"/> while ''W. vanderleueri'' and ''W. alcootaensis'' are estimated to have had bodymasses of {{Convert|30|kg|lb}} and {{Convert|50|kg|lb}}.<ref>{{Cite journal |last1=Warburton |first1=Natalie M. |last2=Yates |first2=Adam. M. |date=2020-12-10 |title=Functional morphology of Wakaleo postcrania from the middle to late Miocene of central Australia reveals new insights in the evolution of marsupial hypercarnivores |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2021.1878203 |journal=Journal of Vertebrate Paleontology |language=en |volume=40 |issue=6 |pages=e1878203 |doi=10.1080/02724634.2021.1878203 |bibcode=2020JVPal..40E8203W |s2cid=233793061 |issn=0272-4634|url-access=subscription }}</ref>

=== ''Wakaleo alcootaensis'' ===
The first description was given in an examination of material discovered at [[Alcoota]] in the [[Northern Territory]] of Australia. The holotype was a single fossil maxilla fragment<ref name="australian_museum_1">{{cite web |title=Australian Museum - Wakaleo |url=http://australianmuseum.net.au/Wakaleo-vanderleuri |accessdate=2011-01-17}}</ref> found in 1974 by the palaeontologist [[Michael Archer (palaeontologist)|Michael Archer]].

Fossil material of this species is fragmentary and rare, and it is only known amongst the Alcoota local fauna. Further evidence of the animals cranial and dental features were examined in 2014, leading to a revision of ''Wakaleo alcootaensis'' that provided further support to the separation from earlier ''Wakaleo'' species.<ref name="Yates2015">{{cite journal |last1=Yates |first1=Adam M. |date=12 November 2015 |title=New craniodental remains of Wakaleo alcootaensis (Diprotodontia: Thylacoleonidae) a carnivorous marsupial from the late Miocene Alcoota Local Fauna of the Northern Territory, Australia |journal=PeerJ |language=en |volume=3 |pages=e1408 |doi=10.7717/peerj.1408 |issn=2167-8359 |pmc=4647553 |pmid=26587359 |doi-access=free }}</ref>

=== ''Wakaleo oldfieldi'' ===
''Wakaleo oldfieldi'' is an [[extinct]] species of [[Thylacoleonidae|marsupial lions]] of the genus ''Wakaleo'', found in [[Miocene]] deposits of [[South Australia]]. It had three unfused molar teeth instead of two fused molars as is the case with the [[Pleistocene]] ''[[Thylacoleo carnifex]]''. As with ''[[Thylacoleo carnifex]]'', this species is presumed to have used its [[maxilla]]ry (upper) teeth to hold its food and sharpen the [[Mandible|mandibular]] teeth, the latter were also used in slicing and stabbing during eating. The [[premolar]]s also had a crescent-shaped circumference for slicing.<ref>{{cite web |title=Thylacoleonid Genera and Species - Tertiary Genera |url=http://www.naturalworlds.org/thylacoleo/genera/tertiary/GS_tertiary_2.htm |access-date=1 October 2022 |website=www.naturalworlds.org}}</ref>

=== ''Wakaleo pitikantensis'' ===
The description by Rauscher was published in 1987, naming a new species and genus as ''Priscileo pitikantensis''.<ref name=":0" /> The designation as the type species of the genus ''[[Priscileo]]'' was later recognised as a species of a revised description of ''Wakaleo''.<ref name="Gillespie2017" /> Fossil material examined by Rauscher was obtained at Lake Pitikanta, situated to the east of [[Lake Eyre]].<ref name=":0" />

The smallest known species of ''Wakaleo'', it lived in Australia about 25 million years ago, from the late [[Oligocene]] to middle [[Miocene]], and was approximately the size of a cat. They were mid-sized predators who probably hunted in trees or ambushed prey from a branch. Like the later discovery, ''[[Wakaleo schouteni]]'', the species possesses three premolars and four molars which distinguishes them from others of the genus. A little smaller than ''W. schouteni'', this species also differs in the morphology of the humerus.<ref name="Gillespie2017" />

It is known only from a few post cranial bones and a poorly preserved maxillary fragment found at the Lake Pitikanta site of the [[Lake Ngapakaldi to Lake Palankarinna Fossil Area]] in northeastern South Australia. To date ''W. pitikantensis'', ''Wakaleo schouteni'' and ''"Priscileo' roskellyae'' are the most ancient phylogenies of the [[Thylacoleonidae]] and presumed to have diverged during the early to middle Oligocene epoch.<ref name="Gillespie2017" />

=== ''Wakaleo schouteni'' ===
The description of the species was published in 2017, the collaborating authors [[Anna Gillespie]], [[Mike Archer (paleontologist)|Mike Archer]] and [[Suzanne Hand]] working on the PANGEA research project based at the University of New South Wales. The [[holotype]] is a largely complete skull retaining some teeth and alveoli, with other materials such as the lower jaw and a humerus associated with the new species as paratypes. The discovery of more complete evidence of Oligocene species of the marsupial lion lineages prompted the authors to emend the circumscription of ''Wakaleo'' to include the type species of another genus, ''[[Priscileo pitikantensis]]'', as a sister species to this taxon and contradicting a 2016 study that supported separation of ''P. pitikantensis'' from the wakaleo clade. Another early [[thylacoleonid]] species, ''[[Priscileo roskellyae|"Priscileo" roskellyae]]'', was determined to have diverged from this genus during early period of the fossil record, and scant evidence of the species did not allow revising authors to assign a taxonomic placement with any confidence.<ref name="Gillespie2017" />

A painter celebrated for his reconstructions of ancient animals, [[Peter Schouten]], was honored by the authors in their proposed epithet for the species.<ref name="Gillespie2017" />

A species assigned to ''Wakaleo'' and resembling a contemporary animal of the Oligocene, ''Wakaleo pitikantensis''. They were mid-sized predators who probably hunted in trees or ambushed prey from a branch. Calculations of the size following a method of statistical analysis for predicting body size, derived from 164 millimetres for the greatest length of the skull, indicate a weight of 22.6 or 24.0 kilograms.<ref name="Gillespie2017" /><ref name="Myers2001">{{cite journal |last1=Myers |first1=T.J. |date=2001 |title=Prediction of marsupial body mass |journal=Australian Journal of Zoology |volume=49 |issue=2 |pages=99 |doi=10.1071/ZO01009}}</ref> Another estimate using [[regression equation]]s previously used in calculations of body size for ''[[Thylacoleo carnifex]]'', a very large carnivore, resulted in a value around 5&nbsp;kg that the describing authors considered too small.<ref name="Wroe2003">{{cite journal |last1=Wroe |first1=Stephen |last2=Myers |first2=Troy |last3=Seebacher |first3=Frank |last4=Kear |first4=Ben |last5=Gillespie |first5=Anna |last6=Crowther |first6=Mathew |last7=Salisbury |first7=Steve |date=September 2003 |title=An alternative method for predicting body mass: the case of the Pleistocene marsupial lion |url=https://www.researchgate.net/publication/52005060 |journal=Paleobiology |volume=29 |issue=3 |pages=403–411 |doi=10.1666/0094-8373(2003)029<0403:AAMFPB>2.0.CO;2 |s2cid=85776991}}</ref> The dentition suggests that ''W. schouteni'' may have been omnivorous, but reveals the transition to hyper-carnivory of the genus during the Miocene.<ref name="Gillespie2017" />

The distribution of the species included the [[Riversleigh World Heritage Area]] in the northwest of Queensland, at sediments dated to the later Oligocene to early Miocene. The assumption of at least partially arboreal habits is by the forest types that existed during the Oligo-Miocene at Riversleigh, open woodlands with the later development of rainforest. The humerus morphology is proposed to support this conception as a scansorial and arboreal predator, being similar to the structures of the arm and shoulder in species such as ''[[Phascolarctos cinereus]]'', the related koala, and spotted cuscus ''[[Spilocuscus maculatus]]''.<ref name="Gillespie2017" />

=== ''Wakaleo vanderleuri'' ===
The type species fossils were originally found in the [[Bullock Creek (Northern Territory)]] area. It has been found in limestone deposits in [[Australian Fossil Mammal Sites (Riversleigh)|Riversleigh]].<ref name="wakaleo.net">{{cite web |title=''Wakaleo vanderleueri'' Vanderleuer's Little Lion |url=http://www.wakaleo.net/?p=744 |url-status=dead |archive-url=https://web.archive.org/web/20150310031739/http://www.wakaleo.net/?p=744 |archive-date=10 March 2015 |accessdate=28 May 2019 |website=Riversleigh faunal encyclopedia |publisher=wakaleo.net}}</ref>

The first fossil, a right dentary fragment, was found in 1967 by a group of field workers at Bullock Creek in the Northern Territory. A left dentary of the species in a good state of preservation was described in 1896.<ref name="Megirian1986">{{cite journal |last1=Megirian |first1=D. |date=28 January 1986 |title=The dentary of ''Wakaleo vanderleueri'' (Thylacoleonidae: Marsupialia) |url=https://www.biodiversitylibrary.org/part/260875 |journal=The Beagle: Occasional Papers of the Northern Territory Museum of Arts and Sciences |volume=3 |pages=71–79}}</ref> Since then more fossils have been recovered, including a well-preserved near-complete cranium with the mandible intact.<ref name="wakaleo.net" />

== Ecology ==
The various species of ''Wakaleo'' are thought to have been carnivores.<ref name="Gillespie2023">{{cite journal |last1=Gillespie |first1=A. K. |date=2023 |title=Two new marsupial lion taxa (Marsupialia, Thylacoleonidae) from the early and Middle Miocene of Australia |journal=Alcheringa: An Australasian Journal of Palaeontology |pages=1–16 |doi=10.1080/03115518.2022.2152096 |s2cid=256157821}}</ref> Later species of ''Wakaleo'' show increased adaptation to terrestrial locomotion (as opposed to the [[Arboreal locomotion|arboreal]] lifestyle of earlier species), and to grappling prey with their forelimbs,<ref>{{Cite journal |last=Warburton |first=Natalie M. |last2=Yates |first2=Adam. M. |date=2020-12-10 |title=Functional morphology of Wakaleo postcrania from the middle to late Miocene of central Australia reveals new insights in the evolution of marsupial hypercarnivores |url=https://www.tandfonline.com/doi/full/10.1080/02724634.2021.1878203 |journal=Journal of Vertebrate Paleontology |language=en |volume=40 |issue=6 |pages=e1878203 |doi=10.1080/02724634.2021.1878203 |issn=0272-4634|url-access=subscription }}</ref> likely correlated with increasing prey size.<ref name="Yates2015" /><ref name="Gillespie2017" />

==References==
{{Reflist}}
*[http://www.naturalworlds.org/thylacoleo/index.htm Tertiary Genera of Thylacoleonidae]
*[http://www.fmnh.helsinki.fi/users/haaramo/Metazoa/Deuterostoma/Chordata/Synapsida/Metatheria/Notometatheria/Diprotodontia/diprotodontia.htm Mikko's Phylogeny Archive]

{{Vombatiformes}}
{{Taxonbar|from=Q140445}}

[[Category:Wakaleo| ]]
[[Category:Miocene mammals of Australia]]
[[Category:Miocene marsupials]]
[[Category:Prehistoric marsupial genera]]
[[Category:Fossil taxa described in 1974]]
[[Category:Taxa named by William A. Clemens Jr.]]
[[Category:Carnivorous marsupials]]