Migrations, diversifications and extinctions: the evolutionary history of crocodyliforms in Australasia

Figures & data

Figure 1. Extant Crocodylus from Australasia. A, Crocodylus johnstoni. B, Crocodylus porosus.

Figure 2. Australasian fossil crocodyliform localities. (1) Busai, Muyua, Papua New Guinea (de Vis 1905, Molnar 1982a); (2) Awe Fauna, Otibanda Formation, Papua New Guinea (Plane 1967); (3) Windjana Gorge, Western Australia (Gorter & Nicoll 1978); (4) Quanbun Local Fauna, Western Australia (Flannery 1984); (5) Bullock Creek, Northern Territory (Willis et al. 1990, Megirian et al. 1991, Murray & Megirian 1992, Megirian 1994, Stein et al. 2017, 2020, Yates 2017); (6) Alcoota Station, Northern Territory (Stirton 1967, Woodburne 1967, Murray & Megirian 1992, Murray et al. 1993, Megirian et al. 1996, Stein et al. 2017, Yates 2017); (7) Kangaroo Well Local Fauna, Ulta Limestone, Northern Territory (Megirian et al. 2004); (8) Pwerte Marnte Marnte Local Fauna, Northern Territory (Murray & Megirian 2006, Yates 2017); (9) Warburton River, South Australia (Willis & Molnar 1997a); (10) Wipajiri Formation, Lake Ngapakaldi, South Australia (Stirton et al. 1961, 1967, Willis & Molnar 1991b); (11) Cooper’s Creek, South Australia (Willis & Molnar 1997a, Vickers-Rich & Rich 1999); (12) Lake Palankarinna, Etadunna Formation (Stirton et al. 1961, Willis & Molnar 1991b), Tirari Formation, and Mampuwordu Sands, South Australia (Hecht & Archer 1977, Yates & Pledge 2017); (13) Lakes Pinpa and Tarkarooloo, Namba Formation, South Australia (Willis & Molnar 1991b). (14) Dinosaur Cove, Otway Ranges, Eumeralla Formation, Victoria (Willis 1997b, Salisbury et al. 2003, Poropat et al. 2018, Paragnani et al. 2019); (15) Krui River Local Fauna, New South Wales (Molnar 1991); (16) Myrtle Vale, New South Wales (Thompson 1980, Willis & Molnar 1997a); (17) Cuddie Springs, New South Wales (Dodson et al. 1993); (18) Griman Creek Formation, Lightning Ridge, New South Wales (Etheridge 1917, Molnar 1980, Molnar & Willis 2001, Hart 2020, Hart et al. 2019, 2021); (19) Texas Caves, Queensland (Hecht & Archer 1977, Archer 1978, Molnar 1982c); (20) Multiple localities on the Darling Downs (King Creek, west of Pilton, Westbrook or Gowrie Creek, Toowoomba Region), Queensland (Lydekker 1888, Molnar 1982c, Sobbe et al. 2013, Ristevski et al. 2020a, 2021); (21) Geebung, Runcorn, Eight Mile Plains and Redbank Plains, Brisbane area, Queensland (Jones 1927, Riek 1952, Willis & Molnar 1991a); (22) Macalister, Queensland (Longman 1928, Molnar 1982b, Willis & Molnar 1997a); (23) Chinchilla, Queensland (de Vis 1886, Hill et al. 1970, Gorter & Nicoll 1978, Molnar 1982b, 1982c, Willis & Molnar 1997a, Mackness et al. 2010, Louys & Price 2015, Chiotakis 2018, Ristevski et al. 2020a, Campbell et al. 2021) and Brigalow, Queensland (Longman 1929); (24) Tingamarra Local Fauna, Murgon, Queensland (Molnar 1982b, Willis et al. 1993, Salisbury & Willis 1996, Stein et al. 2012, 2020); (25) The Narrows Graben, near Gladstone, Queensland (Hills 1943, Coshell 1986, Holt et al. 2005, Buchanan 2008, 2009, Stein et al. 2017); (26) Mt. Etna region, Queensland (Hocknull 2009); (27) Lansdowne Station, near Tambo, Queensland (Longman 1925); (28) Winton Formation, near Isisford, Queensland (Salisbury et al. 2006, Syme & Salisbury 2018); (29) Winton Formation at Elderslie Station (White et al. 2022) and Karoola Station (Poropat et al. 2021); (30) South Walker Creek (Hocknull et al. 2020) and near the Nebo district (Willis & Molnar 1997a), Queensland; (31) Cape Hillsborough, Queensland (McNamara 1993); (32) Bluff Downs Local Fauna, Allingham Formation, Queensland (Archer in Archer & Wade 1976, Molnar 1979, Willis & Mackness 1996, Mackness & Sutton 2000); (33) Wyandotte Local Fauna, Wyandotte Formation, Queensland (McNamara 1990); (34) Tara Creek, near Maryvale, Queensland (Longman 1924, Willis & Molnar 1997a, Molnar 1982c); (35) Riversleigh World Heritage Area, Queensland (Willis & Archer 1990, Willis et al. 1990, Willis 1993b, 1995, 1997a, 2001, Willis & Molnar 1997a, Scanlon 2006a, 2014, Yates 2017, Stein et al. 2015, 2016, 2017, 2020, Ristevski et al. 2023); (36) “Rosella Plains” Station, Queensland (Molnar 1982c); (37) Floraville Downs Station, Queensland (Willis & Archer 1990, Willis & Molnar 1997a, Molnar et al. 2017, Ristevski et al. 2019, Stein et al. 2020); (38) “Alehvale” Station, Croydon area (Molnar 1982c); (39) Tea Tree Cave, Chillagoe area, Queensland (Molnar 1982c); (40) “Glen Garland” Station, Queensland (Molnar 1982c); (41) Bannockburn Formation, South Island, New Zealand (Molnar & Pole 1997, Salisbury et al. 2017); (42) Kanumera, Isle of Pines, New Caledonia (Buffetaut 1983, Balouet & Buffetaut 1987); (43) Pindai Caves, Grande Terre, New Caledonia (Balouet & Buffetaut 1987, Anderson et al. 2010, Salisbury et al. 2010); (44) Arapus and Teouma archaeological sites, Efate, Vanuatu (Mead et al. 2002, Hawkins 2015); (45) Voli Voli Cave, Viti Levu, Fiji (Molnar et al. 2002); (46) Wainibuku Cave, Viti Levu, Fiji (Molnar et al. 2002); (47) Naigani, Fiji (Irwin et al. 2011).

Figure 3. Stratigraphical age ranges of documented Australasian crocodyliform taxa. Red asterisk * denotes the uncertain late Paleocene to early Eocene age of the ‘Runcorn taxon’; Pliocene–Pleistocene Gunggamarandu maunala; mid-Pliocene to ?Late Pleistocene Paludirex vincenti; ?Pleistocene ‘Gavialis papuensis’. Information from the Paleobiology Database (https://paleobiodb.org) accessed on 11 July 2022. Numerical ages from version 2021/10 of the International Chronostratigraphic Chart (https://stratigraphy.org/chart). Abbreviations: Oligo., Oligocene; Paleo., Paleocene; Pleisto., Pleistocene; Plio., Pliocene.

Figure 4. Published studies on Australasian crocodyliforms. A, Peer-reviewed articles exclusively documenting or containing substantial information about Australasian crocodyliforms. Conference abstracts, unpublished theses, and non-peer reviewed articles are excluded. B, Number of formally published peer-reviewed articles dedicated to a single extinct crocodyliform genus from Australasia. *Combined articles dedicated to Paludirex and ‘Pallimnarchus’. Silhouette of Paludirex vincenti in A from PhyloPic by Armin Reindl. For more information see Supplemental Data S1.

Table 1. List of crocodyliform taxa from Australasia (classification based on the results from Fig. 24).

CROCODYLOMORPHA Hay, 1930 (sensu Nesbitt, 2011) CROCODYLIFORMES Hay, 1930 (sensu Sereno et al., 2001)   MESOEUCROCODYLIA Whetstone & Whybrow, 1983 (sensu Sereno et al., 2001)    NEOSUCHIA Benton & Clark, 1988 (sensu Sereno et al., 2001)     †Confractosuchus sauroktonos White et al., 2022     †SUSISUCHIDAE Salisbury et al., 2003      †Isisfordia duncani Salisbury et al., 2006      †Isisfordia molnari Hart et al., 2019      †Isisfordia selaslophensis (Etheridge, 1917) Hart, 2020         =†Crocodilus (?Botosaurus) selaslophensis Etheridge, 1917          =†Crocodylus (Bottosaurus) selaslophensis (Etheridge, 1917) Molnar, 1980     EUSUCHIA Huxley, 1875      CROCODYLIA Gmelin, 1789 (sensu Clark in Benton & Clark, 1988)       LONGIROSTRES Harshman et al., 2003        GAVIALOIDEA Hay, 1930         †‘Gavialis papuensis’ de Vis, 1905         †Gunggamarandu maunala Ristevski et al., 2021         †Harpacochampsa camfieldensis Megirian et al., 1991        CROCODYLOIDEA Fitzinger, 1826 (sensu Brochu, 2003)         †MEKOSUCHINAE Willis et al., 1993          †Australosuchus clarkae Willis & Molnar, 1991b          †Kalthifrons aurivellensis Yates & Pledge, 2017          †Kambara implexidens Salisbury & Willis, 1996          †Kambara molnari Holt et al., 2005          †Kambara murgonensis Willis et al., 1993          †Kambara taraina Buchanan, 2009          †MEKOSUCHINI Salisbury & Willis, 1996           †‘Baru’ huberi Willis, 1997a           †Baru darrowi Willis et al., 1990           †Baru wickeni Willis, 1997a           †Mekosuchus inexpectatus Balouet & Buffetaut, 1987           †Mekosuchus kalpokasi Mead et al., 2002           †Mekosuchus sanderi Willis, 2001           †Mekosuchus whitehunterensis Willis, 1997a           †Paludirex gracilis (Willis & Molnar, 1997a) Ristevski et al., 2020a            =†Pallimnarchus gracilis Willis & Molnar, 1997a           †Paludirex vincenti Ristevski et al., 2020a           †Quinkana babarra Willis & Mackness, 1996           †Quinkana fortirostrum Molnar, 1982c           †Quinkana meboldi Willis, 1997a           †Quinkana timara Megirian, 1994           †Trilophosuchus rackhami Willis, 1993b           †Volia athollandersoni Molnar et al., 2002        CROCODYLIDAE (Cuvier, 1807)         Crocodylus halli Murray et al., 2019         Crocodylus johnstoni (Krefft, 1873)         Crocodylus novaeguineae (Schmidt, 1928)         Crocodylus porosus Schneider, 1801          =Crocodylus nathani Longman, 1924 CROCODYLOMORPH ICHNOTAXA Hatcherichnus ichnosp. indet. Foster & Lockley, 1997 (Poropat et al., 2021)

Revised from Thorn et al. (2021b). Note Isisfordia molnari may be a junior synonym of Isisfordia selaslophensis (see Hart 2020); placement of Mekosuchinae within Longirostres and as a subclade of Crocodyloidea is uncertain; placement of Kalthifrons aurivellensis outside of Mekosuchini is tentative; ‘Baru’ huberi is not referrable to the genus Baru (see Yates 2017, Lee & Yates 2018); Ultrastenos willisi requires revision; ‘Gavialis papuensis’ requires revision; Crocodylus nathani requires revision; taxonomic validity of Crocodylus halli is based on morphometric data.

Table 2. List of all currently named crocodyliform species from Australasia.

Taxon	Age	Locality/Distribution	Source
Australosuchus clarkae	Chattian, late Oligocene	Etadunna Formation, Lake Eyre Basin, SA; Namba Formation, Lake Eyre Basin, SA	Willis & Molnar (1991b); Yates (2017)
‘Baru’ huberi	Chattian, late Oligocene	White Hunter, Riversleigh WHA, QLD	Willis (1997a)
Baru darrowi	Langhian–middle Serravallian, Middle Miocene	Ringtail, Riversleigh WHA, QLD; Bullock Creek, Camfield station, NT	Willis et al. (1990); Yates (2017)
Baru wickeni	Chattian, late Oligocene	D Site and White Hunter, Riversleigh WHA, QLD	Willis (1997a); Yates (2017)
Confractosuchus sauroktonos	Cenomanian–early Turonian, Late Cretaceous	Winton Formation, Elderslie Station, QLD	White et al. (2022)
Crocodylus halli	present (no known fossil record)	southern New Guinea	Murray et al. (2019); extant
Crocodylus johnstoni	Late Pleistocene–present	Terrace Site, Riversleigh WHA, QLD (fossil); 'Leichhardt 3', Floraville Station, QLD (fossil); mainland northern Australia (QLD, NT, WA)	Willis & Archer (1990); Isberg et al. (2017); extant
Crocodylus novaeguineae	present (no known fossil record)	northern New Guinea	Solmu & Manolis (2019); extant
Crocodylus porosus	present (no unambiguous fossil record; see text)	northern Australia, New Guinea, Vanuatu, as well as a vast range in southeast Asia	Webb et al. (2021); extant
‘Gavialis papuensis’	Pleistocene (presumed)	Busai, Muyua, PNG	de Vis (1905); Molnar (1982a)
Gunggamarandu maunala	Pliocene or Pleistocene	Darling Downs, QLD	Ristevski et al. (2021)
Harpacochampsa camfieldensis	Langhian, Middle Miocene	Bullock Creek, Camfield Station, NT	Megirian et al. (1991)
Isisfordia duncani	late Albian, Early Cretaceous	Winton Formation, Isisford, QLD	Salisbury et al. (2006)
Isisfordia molnari	Cenomanian, Late Cretaceous	Griman Creek Formation, Lightning Ridge, NSW	Hart et al. (2019)
Isisfordia selaslophensis	Cenomanian, Late Cretaceous	Griman Creek Formation, Lightning Ridge, NSW	Etheridge (1917); Hart et al. (2019); Hart (2020)
Kalthifrons aurivellensis	Pliocene	Golden Fleece, Lake Palankarinna, SA	Yates & Pledge (2017)
Kambara implexidens	Ypresian, early Eocene	Tingamarra Station, near Murgon, QLD	Salisbury & Willis (1996)
Kambara molnari	middle Eocene–late Eocene	Stuart Deposit, The Narrows Graben, Gladstone QLD	Holt et al. (2005)
Kambara murgonensis	Ypresian, early Eocene	Tingamarra Station, near Murgon, QLD	Willis et al. (1993)
Kambara taraina	middle Eocene–late Eocene	Stuart Deposit, The Narrows Graben, Gladstone QLD	Buchanan (2009)
Mekosuchus inexpectatus	Late Pleistocene–Holocene	Pindaï Caves, Grande Terre, New Caledonia; Kanumera, Ille des Pins, New Caledonia	Balouet & Buffetaut (1987)
Mekosuchus kalpokasi	Holocene	Arapus archaeological Site, Efate Island, Vanuatu	Mead et al. (2002)
Mekosuchus sanderi	late Langhian–middle Serravallian Middle Miocene	Ringtail, Riversleigh WHA, QLD	Willis (2001)
Mekosuchus whitehunterensis	Chattian, late Oligocene	White Hunter, Riversleigh WHA, QLD	Willis (1997a)
Paludirex gracilis	Late Pleistocene	Terrace Site, Riversleigh WHA, QLD	Willis & Molnar (1997a); Ristevski et al. (2020a)
Paludirex vincenti	mid-Pliocene–Pleistocene	Darling Downs and Lansdowne, QLD	Ristevski et al. (2020a); this study
Quinkana babarra	mid-Pliocene	Allingham Formation, Bluff Downs, QLD	Willis & Mackness (1996)
Quinkana fortirostrum	Middle Pleistocene	Tea Tree Cave, near Chillagoe, QLD	Molnar (1982c)
Quinkana meboldi	Chattian, late Oligocene	White Hunter, Riversleigh WHA, QLD	Willis (1997a)
Quinkana timara	Langhian, Middle Miocene	Bullock Creek, Camfield Station, NT	Megirian (1994)
Trilophosuchus rackhami	late Langhian–middle Serravallian, Middle Miocene	Ringtail, Riversleigh WHA, QLD	Willis (1993b); Ristevski et al. (2023)
Ultrastenos willisi	Chattian, late Oligocene	Low Lion, Riversleigh WHA, QLD	Stein et al. (2016)
Volia athollandersoni	Late Pleistocene–Holocene	Voli Voli Cave and Wainibuku Cave, Viti Levu, Fiji	Molnar et al. (2002)

Junior synonyms or nomina dubia (Crocodylus nathani, Pallimnarchus) are not listed. Abbreviations: NSW, New South Wales; NT, Northern Territory; PNG, Papua New Guinea; QLD, Queensland; SA, South Australia; WA, Western Australia; WHA, World Heritage Area.

Figure 5. Confractosuchus sauroktonos, AODF0890, holotype skull in left lateral view.

Figure 6. Isisfordia. A, Isisfordia duncani, QMF36211, holotype, skeleton in dorsal view. B, Isisfordia duncani, QMF44320, skull in dorsal view. C, Isisfordia molnari, AM F125553, holotype, partial cranium in dorsal view. D, Isisfordia selaslophensis, AM F15818, holotype, right maxillary fragment in lateral view. Arrows in C and D indicate anterior.

Figure 7. Australosuchus clarkae, QMF16788, holotype in dorsal view.

Figure 8. Baru. A, Baru wickeni, NTM P91171-1, skull in dorsal view. B, Baru wickeni, NTM P91171-1, skull in left lateral view. C, Baru darrowi, NTM P8695-8, holotype, partial skull in dorsal view. D, Baru darrowi, NTM P8695-8, holotype, partial skull in right lateral view.

Figure 9. ‘Baru’ huberi. QMF31060, holotype. A, Partial snout in dorsal view. B, Partial snout in ventral view.

Figure 10. Kalthifrons aurivellensis, SAMA P35062, holotype, skull in dorsal view.

Figure 11. Kambara. A, Kambara implexidens, QMF29662, holotype, skull in dorsal view. B, Kambara molnari, QMF12364, holotype, left mandible in lateral view.

Figure 12. Mekosuchus from the Riversleigh World Heritage Area. A, Mekosuchus sanderi, QMF31188, holotype, partial left maxilla in lateral view. B, Mekosuchus whitehunterensis, QMF31051, holotype, partial right maxilla in ventral view. C, Mekosuchus sanderi, QMF31166, partial cranium in dorsal view. D, Mekosuchus whitehunterensis, QMF31052, partial frontal in dorsal view. E, Mekosuchus whitehunterensis, QMF31053, right mandible in lateral view. Arrows in A, B, and D indicate anterior. Abbreviation: max om, orbital margin of the maxilla.

Figure 13. Mekosuchus inexpectatus and Volia athollandersoni. A, Mekosuchus inexpectatus, NCP 06, holotype, right mandible in lateral view. B, Volia athollandersoni, NMNZ S37341, holotype (frontal fragment), and NMNZ S37342 (parietal) in dorsal views. C, V. athollandersoni, NMNZ S37332, partial left mandible in lateral view.

Figure 14. Paludirex vincenti. A, ‘Geoff Vincent’s specimen’ (CMC2019-010 + QMF59017), holotype, partial skull, all skull pieces in dorsal view; arrow indicates basicranium (QMF59017) ventral to the cranial table (CMC2019-010-5). B, Skull of ‘Geoff Vincent’s specimen’ in dorsal view; dashed lines indicate hypothetical outline of the skull. QMF1752, digital model of the partial skull in C, dorsal, D, left lateral, E, right lateral, and F, ventral views. A and B are modified from Ristevski et al. (2020a).

Figure 15. Quinkana. A, Quinkana meboldi, QMF31056, holotype, left maxilla in lateral view. B, Quinkana timara, NTM P895-19, holotype, partial snout in right lateral view. C, Quinkana babarra, QMF23220, holotype, right maxillary piece in lateral view. D, Quinkana fortirostrum, AM F57844, holotype, snout in left lateral view. Arrows indicate anterior.

Figure 16. Trilophosuchus rackhami, QMF16856, holotype. A, Cranium in dorsal view. B, Cranium in ventral view.

Figure 17. Skulls of extant Crocodylus from Australasia. A, Crocodylus novaeguineae, QMJ5332, skull in dorsal view. B, Crocodylus halli, UF 145927, digital model of the skull in dorsal view (downloaded from MorphoSource https://www.morphosource.org/concern/media/000039626). C, Crocodylus johnstoni, QMJ58446, skull in dorsal view. D, Crocodylus porosus, QMJ48127, skull in dorsal view.

Figure 18. A, Crocodylus sp. QMF9229, partial rostrum from the Pliocene Allingham Formation, Bluff Downs Queensland. B, Reconstructed rostrum of QMF9229. C, Rostrum of equivalent-sized juvenile Crocodylus porosus (UQSSAL unregistered).

Figure 19. ‘Gavialis papuensis’. A, B, C, QMF406, mandibular fragments in dorsal views. D, Isolated tooth crown, unnumbered QMF specimen. E, F, QMF340, partial cervical vertebrae in right lateral views. G, QMF340, partial dorsal vertebra in left lateral view. H, QMF340, partial sacral vertebra (tentatively interpreted as a second sacral) in right lateral view. I, QMF340, partial caudal vertebra in right lateral view. J, QMF341, isolated osteoderm in dorsal view. K, QMF341, isolated osteoderm in dorsal view. Note that the osteoderm in K has a thin-section shown in L.

Figure 20. Gunggamarandu maunala. QMF548 (=QMF14.548, old registry number), holotype. A, Cranium in dorsal view. B, Cranium in posterior view. C, Hypothetical outline of the skull in dorsal view, with QMF548 depicted in its corresponding position (modified from Ristevski et al. 2021). Skull outline in C is based on skulls of Dollosuchoides densmorei and Kentisuchus spenceri.

Figure 21. Harpacochampsa camfieldensis. A, Holotype cranium, NTM P87106-1, in dorsal view. B, Holotype cranium, NTM P87106-1, in posterior view. C, Partial snout, NTM P87106-5, in ventral view.

Figure 22. ‘Runcorn taxon’. A, QMF73195 (formerly UQF73195), anterior dentary piece in dorsal view. B, QMF73195, reconstructed anterior mandibular portion in dorsal view. C, QMF12363 (formerly UQF12363), cervical centrum in left lateral view. Isolated crocodylian tooth associated with QMF73195 in lingual view; D, scaled to the dentary fragment and centrum; E, scaled for visualization.

Figure 23. ‘Darling Downs taxon’ (= Paludirex? sp. nov.). QMF1154, left premaxilla and fragment of maxilla in A, dorsal and B, ventral views. C, QMF1154, premaxilla in anterior view, with the indicated mirrored photograph in order to aid comparison (modified from Ristevski et al. 2020a). the curved red line in C indicates the anterior arching of the premaxilla. D, QMF1151, incomplete left premaxilla in ventral view. NHMUK PV OR 43047a, right premaxilla and fragment of maxilla in E, ventral and F, dorsal views.

Figure 24. Strict consensus of a single fully resolved most parsimonious cladogram from the analysis run under TrS, and a weighting strength of k = 25. See Supplemental Data S2 for additional information. Abbreviations: CI, consistency index; #MPCs, number of most parsimonious cladograms; #OTUs, number of operational taxonomic units; RI, retention index.

Figure 25. Strict consensus of 129600 MPCs from the analysis run under TrS, without IW. The results from this analysis point towards an alternative position for Mekosuchinae than that recovered in other analyses. Australasian mekosuchines are highlighted in red. See Supplemental Data S2 for additional information. Abbreviations: CI, consistency index; #MPCs, number of most parsimonious cladograms; #OTUs, number of operational taxonomic units; RI, retention index.

Figure 26. 50% majority-rule consensus topology from the analysis run under TrS, without IW. Australasian mekosuchines are highlighted in red. See Fig. 25 for the strict consensus topology of the same analysis. See Supplemental Data S2 for additional information. Abbreviations: CI, consistency index; #MPCs, number of most parsimonious cladograms; #OTUs, number of operational taxonomic units; RI, retention index.

Table 3. Summary of the phylogenetic results.

Search option	Weighting	k value (if IW)	Steps	#MPCs	CI	RI
Traditional	EW	–	1865	129600	0.195	0.687
Traditional	IW	5	112.54302	1	0.191	0.679
Traditional	IW	12	71.07717	2	0.193	0.683
Traditional	IW	25	43.21850	1	0.194	0.686
New Technology	EW	–	1864	182	0.195	0.688
New Technology	IW	5	112.54248	1	0.191	0.679
New Technology	IW	12	71.07490	2	0.193	0.683
New Technology	IW	25	43.19272	1	0.195	0.686

Abbreviations: CI, consistency index; EW, equal weighting; IW, implied weighting; #MPCs, number of most parsimonious cladograms; RI, retention index.

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