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Small fossil vertebrates from Victoria Cave, Naracoorte, South Australia. IV. Reptiles
... Australia is characterized by an exceptional herpetofaunal diversity (e.g., Pianka, 1989) and Pleistocene and Holocene fossil deposits, comprising herpetofaunal remains, are numerous throughout the continent (e.g., Lundelius, 1983;Bourne, 2000, 2009). However, the potential of Australian reptile and amphibian fossils for examining faunal change during this period has widely been neglected, presumably for the same reasons mentioned above (for exceptions see Smith, 1976Smith, , 1982Hope et al., 1977;Pledge, 1990;Price and Sobbe, 2005;Fraser and Wells, 2006;Hocknull et al., 2007;Hollenshead et al., 2011). Additionally, Australian paleoherpetologists are dealing with higher species diversity within fewer subfamilies in any single deposit; when family-or subfamily-level osteology-based identifications are more likely than genus or species level (Villa et al., 2017). ...
... A major difference arising from the comparison with the extant fauna is an apparent absence of gecko and pygopod fossils in the MDC deposit. An absence of geckos and pygopods has further been reported for Victoria Cave (Naracoorte, South Australia; Smith, 1976) and the Seton Rock Shelter deposit (Kangaroo Island;Smith, 1982). Today, geckos or pygopods can be found in the vicinity of all these deposits (Owens and Graham, 2009;Cogger, 2014;Robertson and Coventry, 2019). ...
... Overall, the results of our analyses suggest that changes in aridity and temperature over the last 14,000 years influenced relative abundances and distributions of reptile taxa in southeastern Australia, leading to gradual changes in community composition during this period. These findings add to the results of the few existing studies on changes of small vertebrate species diversity in southeastern Australia during the Quaternary (Wakefield, 1967(Wakefield, , 1969(Wakefield, , 1972Flood, 1973;Smith, 1976Smith, , 1982Hope et al., 1977;Baird, 1992), which we briefly discuss below. ...
The Quaternary Period is characterized by dramatic global climatic changes. Quaternary fossil deposits, which can offer excellent stratigraphic resolution, provide a unique opportunity to understand how fauna respond to past environmental change. Here, we test if the herpetofauna of McEachern’s Deathtrap Cave, a late Pleistocene to Holocene pitfall trap deposit from Victoria, Australia, shows climate-related shifts in taxonomic relative abundance through time. During the last 14,000 years, southeastern Australia experienced pronounced periods of aridity, while temperatures remained relatively stable. We show that the stratigraphic layers of this deposit are characterized by different relative abundances of reptile subfamilies, and that changes in subfamily abundance between layers correlate with known shifts to aridity, based on the percentage of C4 grasses present in the region. We further identify 13 lizard morphotypes from the fossil deposit and compare this diversity with the present-day lizard fauna. Our analyses indicate that gradual changes in community structure, which are typically observed in southeastern Australian vertebrate communities during the Pleistocene–Holocene transition, can partly be explained by changing aridity. These findings represent an important contribution to understanding Quaternary community change in Australia, particularly because evidence of faunal succession of reptile and amphibian communities in Victoria is lacking. Our results further demonstrate the utility of the Australian herpetofaunal fossil record for detecting community responses to past climate change on relatively shallow timescales and at higher levels of taxonomic identification.
... An unidentified chelid is present in the Miocene Bullock Creek Local Fauna of the Northern Territory according to Murray and Megirian (1992), who also recognize an unidentified emydid from the younger Alcoota Local Fauna. Smith, 1976, SAM P16166 (one of the paratypes), Victoria Cave, Naracoorte. These were "giant" snakes, in the sense that the Anaconda and larger pythons are today, commonly reaching 5 or 6 m. ...
... Previously unrecognized features, such as a sagittal crest and low ornament on the dorsal surface, are evident in these specimens. Tedford and Wells (1990) Pledge (1990), following identifications given by Smith (1976), provides the following list of lizards from the Pleistocene Upper Fossil Fauna of Henschke Fossil Cave near Naracoorte, South Australia; Amphibolurus spp., Tiliqua rugosa, Scincid undet. and Varanus sp. ...
... Barrie (1990) provides descriptions of new material of the giant "madtsoine boid" (madtsoiid) snake Wonambi naracoortensis including many skull elements. Previously, this species was known from only eight vertebrae and a jaw fragment (Smith 1976), so Barrie's material representing two individuals and preserving most major skull elements and the postcranial skeleton represents a significant improvement in our knowledge of this species and, indeed, of madtsoiid snakes in general. The Pleistocene age of the Naracoorte Wonambi make this the youngest madtsoiid and suggests that the predominantly Gondwanan madtsoiids had a refuge in Australia for the latter half of the Tertiary. ...
... Diagnosis-Madtsoiid snake with the following unique combination of characters (autapomorphies denoted by asterisks): anteroposteriorly long and dorsoventrally short neural spine (distinct from Alamitophis [Albino, 1986], Dinilysia [Hecht, 1982], Madtsoia [e.g., Simpson, 1933;Mohabey et al., 2011], Najash [Zaher et al., 2009], Nanowana [Scanlon, 1997], Rionegrophis [Albino, 1986], Wonambi [Smith, 1976], and Yurlunggur [Scanlon, 1992]) with posterodorsal expansion (distinct from Herensugea [Rage, 1996] and Menarana [LaDuke et al., 2010]); shallow fossae lateral to neural spine (distinct from Dinilysia, Najash, and all Madtsoiidae, excepting Madtsoia pisdurensis and Sanajeh [Wilson et al., 2010]); zygantrum more gracile than in Gigantophis (Andrews, 1906), Madtsoia, Menarana, Wonambi, and Yurlunggur; neural canal roughly trapezoidal in shape, contrasting with Rionegrophis; depressed cotyle and condyle (distinct from all Madtsoiidae, except in posterior trunk vertebrae of Herensugea and Patagoniophis australiensis [see Scanlon, 2005]); dorsoventrally tall and laterally projecting synapophyses; synapophyses in posterior trunk vertebrae with dorsal margin that extends far dorsally relative to floor of neural canal*; and subcentral fossae exposed laterally*. ...
... By contrast, most madtsoiid trunk vertebrae bear neural spines that maintain similar heights throughout their anteroposterior lengths (Simpson, 1933;Scanlon, 1997;LaDuke et al., 2010). As in other known madtsoiids, the posterior-most portion of the spine overhangs the zygantrum (e.g., Smith, 1976;Scanlon, 1992Scanlon, , 1997Scanlon, , 2005Rage, 1998;LaDuke et al., 2010). The bilateral fossae that sit alongside the neural spine are also found in specimens of Sanajeh et al., 2010) and those referred to the holotype of Najash (Apestegu ıa and Zaher, 2006;Zaher et al., 2009; but see also Palci et al., 2013). ...
... The posterior portion of the keel is smoothly textured, in contrast to the rugose hemal process in Madtsoia spp. (Simpson, 1933;LaDuke et al., 2010;Mohabey et al., 2011), Gigantophis (Andrews, 1906), and Wonambi (Smith, 1976). Such a process is absent in Menarana (Rage, 1999;LaDuke et al., 2010), Nanowana (Scanlon, 1997), and P. australiensis and Alamitophis tingamarra (Scanlon, 1993(Scanlon, , 2005, and in vertebrae referred to the holotype of Najash (Apestegu ıa and Zaher, 2006;Zaher et al., 2009; but see also Palci et al., 2013). ...
We describe three new fossil snakes on the basis of recently discovered vertebrae collected from the Upper Cretaceous (Maastrichtian) Maevarano Formation of Madagascar. One represents a new genus and species of madtsoiid, Adinophis fisaka, the third member of this family recognized from the Maevarano Formation. It exhibits dorsoventrally compressed centra and dorsally placed synapophyses. The second taxon is a new species of the nigerophiid genus Indophis, I. fanambinana, representing the second nigerophiid taxon known from the Maevarano Formation. It shares numerous features with the Indian nigerophiid I. sahnii, including small synapophyses positioned ventrolaterally on distinct synapophyseal processes and a unique cotylar shape in posterior trunk vertebrae. The discovery of I. fanambinana demonstrates a strong biogeographic link with penecontemporaneous snake faunas from India. A third new taxon is represented by a partial centrum that cannot be assessed thoroughly due to its incompleteness; this specimen is conservatively assigned to Serpentes incertae sedis, gen. et sp. indeterminate. It is distinct from other snakes known from the Late Cretaceous of Madagascar in possessing a strongly depressed condyle and relatively large lateral foramina. These new discoveries, together with previous descriptions of snakes from the Maevarano Formation, make the Maevarano snake fauna one of the most taxonomically diverse snake assemblages known from the Mesozoic and the most diverse in terms of body size range.
... A second specimen was located by Paul Willis in the collections of the Department of Geology and Geophysics, University of Sydney, in a batch of Wellington material which appears to have been collected prior to 1915, most probably in Gaden Cave, and later set aside from the rest of the collection by Mahoney (Willis et at., 1992). Mter comparison with the published descriptions of Wonambi naracoortensis Smith, 1976 and other madtsoiids, vertebrae of Wonambi from N aracoorte, and Tertiary material of related taxa from several northern Australian sites, there seems no reason to distinguish the Wellington specimens taxonomically from W naracoortensis. FAMILY MADTSOIIDAE HOFFSTETTER, 1961 WONAMBI NARACOORTENSIS SMITH, 1976 The first vertebra has been registered as F72999 in the AM palaeontological collection. ...
... 1). A weak but distinct narrow keel is present in the midline, strongest in the middle of the vertebra, between the subcentral foramina; these are Table 1 gives linear measurements of both vertebrae, which allow comparison of proportions with the type series (Smith, 1976). Pledge (1992) records about twenty vertebrae and some jaw fragments of large madtsoiid snakes (referred to Wonambi sp. ...
... Other madtsoiids are now also known from a number of Tertiary faunas in northern Australia (Yurlunggur camfieldensis Scanlon, 1992 from Bullock Creek, N.T.; species of Alamitophis and Patagoniophis from Murgon, Qld; and other species of Yurlunggurand Wonambifrom local faunas at Riversleigh, Qld.; Scanlon, 1992, 1993 andin prep.) . The present record brings the number oflocalities for W naracoortensis to five, all in southern Australia: Victoria Cave and Henschke's Cave, Naracoorte, southern South Australia (Smith, 1976;Barrie, 1990); Mammoth Cave and Koala Cave, southwestern Western Australia (fide Molnar, 1982); and now Wellington Caves, eastern New South Wales. These are all Quaternary deposits in limestone cave systems, mostly Late Pleistocene in age but with older and younger remains also present. ...
Two trunk vertebrae of a species of Wonambi (Serpentes, Madtsoiidae) from unknown deposits in the Wellington Caves system are the only specimens of this taxon known from New South Wales, and considered to be Pleistocene in age. The specimens are referred to the type species, W naracoartensis Smith, 1976. Together with other known occurrences of the genus (in South Australia and Western Australia), this record supports an hypothesis of association with temperate climates during the Quaternary. Other speci-mens previously referred to Wonambi are reassessed with implications for the distribution of Wonambiand another large madtsoiid, Yurlunggur Scanlon, 1992.
... Aplin et al. (1999) aligned specimens from late Quaternary cave deposits in Irian Jaya, New Guinea , to the extant taxon Aepypodius arfakianus. The other living taxon reported as a fossil is Leipoa ocellata, which was listed among late Pleistocene remains from the Victoria Fossil Cave, South Australia (van Tets 1974; van Tets & Smith 1974). The earliest record of the family is Ngawupodius minya, recovered from the Late Oligocene Etadunna Formation of northeastern South Australia (Boles & Ivison 1999; Fig. 1); this form was notable for its diminutive size. ...
... The second could be compromised by the apparent co-occurrence of Progura and Leipoa ocellata from Victoria Fossil Cave. This record of Leipoa, identified by van Tets (1974) and van Tets & Smith (1974), consists of fragments of a juvenile skull; no reasons were given for this taxonomic allocation, although at the time of that study, skull material of Progura was not known. Also among the fossils recovered from this site were post-cranial elements of a very small juvenile megapode that could not be referred to any taxon within the family. ...
... The consensus palaeoenvironmental reconstruction is open forest and/or woodland, with grassy understorey and/or open areas, with some heath and wetlands/swamps (Wells et al. 1984, Pledge 1990, Barrie 1997, Brown & Wells 2000). The extant taxa in these de-posits – including birds at Victoria Fossil Cave (van Tets & Smith 1974) – agree with this habitat interpretation. There is nothing in the reconstructed palaeohabitat of the Naracoorte region that is inconsistent with the occurrence of Leipoa ocellata. ...
... Both M. pisdurensis and M. madagascariensis possess prezygapophyses that extend as far or farther laterally than the synapophyses, which is the plesiomorphic character state for madtsoiids (e.g., Scanlon, 2006). Madtsoia bai shares the derived condition, laterally extended synapophyses, with Wonambi, Gigantophis, Yurlunggur, Sanajeh, and possibly M. camposi (Hoffstetter, 1961b;Smith, 1976;Rage, 1998;Scanlon and Lee, 2000;Wilson et al., 2010). The triangular process on the hemal keel is unique to M. pisdurensis, and both M. bai and M. madagascariensis share bifid hemal processes with Wonambi and Yurlunggur. ...
... The triangular process on the hemal keel is unique to M. pisdurensis, and both M. bai and M. madagascariensis share bifid hemal processes with Wonambi and Yurlunggur. Madtsoia pisdurensis possesses well-developed, deep parazygantral foramina, and shares an anteroposterioly narrow and posteriorly angled spinous process with M. bai (Simpson, 1933) and Wonambi naracoortensis (Smith, 1976). It also shares large body size with M. bai, M. madagascariensis, and M. camposi (Simpson, 1933;Hoffstetter, 1961a;Rage, 1998), Yurlunggur (Scanlon, 1992(Scanlon, , 2006, and Wonambi (Smith, 1976;Scanlon and Lee, 2000). ...
... Madtsoia pisdurensis possesses well-developed, deep parazygantral foramina, and shares an anteroposterioly narrow and posteriorly angled spinous process with M. bai (Simpson, 1933) and Wonambi naracoortensis (Smith, 1976). It also shares large body size with M. bai, M. madagascariensis, and M. camposi (Simpson, 1933;Hoffstetter, 1961a;Rage, 1998), Yurlunggur (Scanlon, 1992(Scanlon, , 2006, and Wonambi (Smith, 1976;Scanlon and Lee, 2000). The extremely large body size of Gigantophis (precondylar centrum length = 3.06 cm; CGM 10022) exceeds estimates for all other madtsoiids (Head and Polly, 2004) and is here considered an autapomorphy of the genus. ...
We report the discovery of a new species of the snake Madtsoia from infratrappean horizons of Late Cretaceous age in Pisdura, central India. Recovered vertebrae are large (1.83 cm long; 4.35 cm tall) and pertain to a snake that was ca. 5 m long. Discovery of Madtsoia in India extends the geographic distribution of the genus and represents only the second species known from the Cretaceous. Vertebrae of Madtsoia pisdurensis sp. nov. are strikingly similar to those of M. bai and M. camposi (South America) and M. madagascariensis (Madagascar), but can be distinguished from them by a unique process on the hemal keel, which is low, flat, and triangular in outline. Whereas the eastern Gondwanan species of Madtsoia (M. madagascariensis, M. pisdurensis) are Late Cretaceous in age, the western Gondwanan species (M. bai, M. camposi) are Paleogene in age. Geophysical evidence suggests that land connections between South America, Madagascar, and Indo-Pakistan were severed by at least 100–90 Ma, which implies that Madtsoia achieved its broad geographic distribution either by (1) origin and dispersion before the end of the Turonian; or (2) the presence of an unrecognized land connection persisting into the latest Cretaceous. Both hypotheses predict that Madtsoia will be discovered in Mesozoic strata of South America, where it survived the Cretaceous-Paleogene mass extinction.
... Descriptions of some of this material have appeared (Barrie 1990;Scanlon 1997;Lee 2000, 2002;Scanlon 2003) but the evidence for a relatively basal phylogenetic position of Madtsoiidae among snakes, and unusually good preservation of the Australian material, justify more detailed documentation. The description of Wonambi naracoortensis bySmith (1976)was the first report of an extinct snake species in Aus− tralia. When first described this species was represented by eight large vertebrae and a jaw fragment from the Main Fos− sil Chamber, Victoria Fossil Cave, Naracoorte, south−eastern South Australia; Smith referred Wonambi to Boidae (as diag− nosed byHoffstetter and Gasc 1969) and considered it to have probable affinities to Madtsoia and Gigantophis, al− though she did not formally include it in Madtsoiinae (erected for the latter two genera by Hoffstetter 1961; spelt incorrectly by Smith 1976 as " Madstoiinae " ). ...
... The description below is based on all known cranial material of Wonambi naracoortensis, which comes from four localities (see previous section), all in South Australia: VF (Pleistocene;Smith 1976;Reed and Bourne 2000), HQ (Pleistocene;Barrie 1990Barrie , 1997), CL (?earlyPliocene;Pledge 1992), and CQ (fis− sure RF 95, probably Pleistocene based on associated mam− mals, James McNamara, personal communication 2002). All of this material is registered in the SAM P or FU collections. ...
... (A) When Wonambi was represented by just a handful of vertebrae, it was referred to Madtsoiidae provisionally or doubtfully, because its parazygantral foramina (not in all ver− tebrae, but in some including the holotype; Smith 1976:fig. 2E) were smaller or less distinct than in the few giant species of Madtsoia and Gigantophis then known (Smith 1976;Rage 1984). According toRieppel et al. (2002: 812), " the associa− tion of Wonambi with madtsoiids was considered tentative by Rage (1984; see also Smith 1976), and it still is. ...
New material and additional morphological details of a rare and phylogenetically significant large fossil snake, Wonambi naracoortensis Smith, 1976, are described from Pleistocene and Pliocene cave deposits in southern South Australia. The new data refute some previous interpretations of the morphology of this species, and have implications for the phylogenetic position of Wonambi relative to extant snakes and other fossils, including other Madtsoiidae. The nature of contacts among palatal, braincase, snout, and mandibular elements imply similar functional attributes to those of extant anilioid snakes: maxillae from multiple individuals show corrugated contact surfaces for the prefrontal, implying a tight suture; structures on the anterior and medial surfaces of the palatine choanal process are interpreted as forming extensive contacts with the vomer and parasphenoid; and the distinctly bounded facets on the basipterygoid processes and pterygoid imply little or no capacity for anteroposterior sliding of the palatopterygoid arch, hence absence of the macrostomatan “pterygoid walk”. On the frontal, interolfactory pillars were either absent or very slender, and a deep, sculptured contact surface for the nasal implies a prokinetic joint was also absent. Margins of the frontal and parietal indicate broad entry of the sphenoid into the ophthalmic fenestra, as in Dinilysia. Similarity of elements and features of the braincase (trigeminal foramen, ear region, and basipterygoid processes) with both lizards and extant snakes show that differences between snakes and other squamates have sometimes been overstated. The case for macrostomatan affinities of Wonambi is not supported by new evidence.
... These records show that distributions of some species were once more extensive than at present. Gigantism characterized a few taxa, such as the giant coucal Centropus colossus (Baird, 1985) and megapode Progura gallinacea (van Tets, 1974); current work by the author indicates that P. gallinacea is actually the megafaunal form of the living malleefowl (Leipoa ocellata ). Flamingos and palaelodids persisted to this time. ...
... Some have well studied records that are temporally restricted and thus not extensive enough to allow useful extrapolation beyond those particular fossils; others with longer records have had only a portion examined in detail. These groups include the Tertiary pelicans (Miller, 1966; Rich and van Tets, 1981), Quaternary rails (Olson, 1975 ) and waterfowl (Olson, 1977), and giant megapodes Progura (van Tets, 1974). A few families have diverse, well-documented records, which provide information on the radiation and diversity of the groups—from the late Oligocene to the present; however, critical periods before this remain unknown. ...
... Comparisons indicate that the new Indian vertebrae do not belong to the genera Wonambi (Smith, 1976) or Yurlunggur (Scanlon, 2006). Wonambi, from the Pliocene and Pleistocene of Australia, presents a high postero-dorsally oriented neural spine running from the anteriormost to the posteriormost extremity of the neural arch, the zygosphene is as wide as high, the zygapophyses are inclined 25° above the horizontal, and a haemal keel is present (Smith, 1976;Scanlon and Lee, 2000). ...
... Comparisons indicate that the new Indian vertebrae do not belong to the genera Wonambi (Smith, 1976) or Yurlunggur (Scanlon, 2006). Wonambi, from the Pliocene and Pleistocene of Australia, presents a high postero-dorsally oriented neural spine running from the anteriormost to the posteriormost extremity of the neural arch, the zygosphene is as wide as high, the zygapophyses are inclined 25° above the horizontal, and a haemal keel is present (Smith, 1976;Scanlon and Lee, 2000). The haemal keel of Wonambi ends with two protuberances, as in Madtsoia and the Tadkeshwar specimens. ...
The Ypresian Cambay Shale Formation at Vastan and Mangrol lignite mines in Gujarat, western India, has yielded a rich vertebrate fauna with numerous taxa of European affinities. Here we report a new, approximately contemporary vertebrate assemblage from two fossiliferous layers in the nearby mine of Tadkeshwar. These layers have yielded a similar mammal fauna with the co-occurrence of the perissodactyl-like cambaytheriid Cambaytherium thewissi, the adapoid primates Marcgodinotius indicus and cf. Asiadapis cambayensis, and the hyaenodontid Indohyaenodon raoi. The presence of these species in both Vastan and Tadkeshwar mines and at different levels suggests that the deposits between the two major lignite seams represent a single land mammal age. Apart from the aforementioned species there is a new, smaller species of Cambaytherium, and a new genus and species of esthonychid tillodont. This fauna also contains the first large early Eocene vertebrates from India, including an unidentified Coryphodon-like pantodont, a dyrosaurid crocodyliform and a new giant madtsoiid snake. Among the Tadkeshwar vertebrates several taxa are of Gondwana affinities, such as Pelomedusoides turtles, dyrosaurids, and large madtsoiids, attesting that the early Eocene was a crucial period in India during which Laurasian taxa of European affinities co-existed with relict taxa from Gondwana before the India-Asia collision. Our results suggest that terrestrial faunas could have dispersed to or from Europe during episodes of contact between the Indian subcontinent and different island blocks along the northern margin of the Neotethys, such as the Kohistan–Ladakh island-arc system. Gondwana taxa might represent remnants of ghost lineages shared with Madagascar, which reached the Indian subcontinent during the late Cretaceous; alternatively they might have come from North Africa and passed along the southern margin of the Neotethys to reach the Indian subcontinent. These dispersals would have been possible as a result of favourable paleogeographic conditions such as the particular Neotethys conformation during the beginning of the early Eocene.
... Comparisons with the different madtsoiid taxa throughout the text were made using published descriptions, as follows: Alamitophis argentinus (Albino, 1986(Albino, , 1987(Albino, , 2000, A. elongatus (Albino, 1994), A. tingamarra (Scanlon, 1993(Scanlon, , 2005a, Gigantophis garstini (Andrews, 1901), Herensugea caristiorum (Rage, 1996(Rage, , 1999, Madtsoia bai (Simpson, 1933), Ma. camposi (Rage, 1998), Ma. madagascariensis (Hoffstetter, 1961;LaDuke et al., 2010), Ma. pisdurensis (Mohabey et al., 2011), Menarana laurasiae (Rage, 1996, 1999), Me. nosymena (LaDuke et al., 2010, Nanowana godthelpi and Na. schrenki (Scanlon, 1997), Patagoniophis parvus (Albino, 1986), P. australiensis (Scanlon, 1993(Scanlon, , 2005a, Rionegrophis madtsoioides (Albino, 1986), Sanajeh indicus , Wonambi naracoortensis (Smith, 1976;Scanlon, 2005b), W. barriei (Scanlon and Lee, 2000), and Yurlunggur camfieldensis (Scanlon, 1992(Scanlon, , 2004; unless otherwise noted, comparisons with these taxa are based on the above references. ...
... Nevertheless, their distribution is highly skewed towards the southern continental fragments of Gondwana that yielded the largest part of the currently known madtsoiids; besides the already listed European occurrences, their scanty northern, Laurasian record was recently supplemented by the report of a possible madtsoiid in Upper Cretaceous beds of North America (see Longrich et al., 2012 (Fig. 6). A southern distribution of the madtsoiids characterizes the Cenozoic as well, when this clade was apparently present in most major southern landmasses such as India (Rage et al., 2008), Africa (Andrews, 1901;Augé and Rage, 2006), South America (Simpson, 1933;Rage, 1998), and most notably Australia (e.g., Scanlon, 1992Scanlon, , 1993Scanlon, , 1997, where it survived into the Pleistocene (Smith, 1976;Scanlon and Lee, 2000). ...
Madtsoiidae are a group of archaic snakes, widely distributed in the Upper Cretaceous of Gondwanan landmasses (South America, Madagascar, India, Africa), but otherwise reported outside Gondwana based only on scarce material from a few southern European localities. Here, we describe associated snake remains from uppermost Cretaceous (Maastrichtian) beds of the Haţeg Basin, Romania, as a new small-madtsoiid taxon, Nidophis insularis, gen. et sp. nov. Nidophis, represented by a large number of well-preserved vertebrae and ribs that apparently belonged to one individual, is one of the best-known Cretaceous madtsoiids, and the most completely documented member of the family from Europe. Phylogenetic analyses place the new taxon within a moderately supported Madtsoiidae, closely related to Herensugea from the Upper Cretaceous of Spain. The two European taxa, together with other small madtsoiids, are recovered as the sister taxon to a second madtsoiid clade including large-to-gigantic forms such as Madtsoia, Wonambi, and Yurlunggur. The presence of these small madtsoiids, together with that of Menarana, in the uppermost Cretaceous of Europe, suggests that early widespread distribution of madtsoiids, extending over the southern part of Europe, instead of late northward immigration from Gondwanan landmasses, as proposed previously, might account for the paleobiogeographic distribution of Cretaceous Madstoiidae.
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... However, the relatively few described Australian Quaternary squamate reptiles (lizards and snakes) largely mirror extant morphologies [5,6]. Two notable exceptions, the giant monitor lizard Varanus (Megalania) priscus [7] and primitive snake Wonambi naracoortensis [8], suggest that any appearance of stable squamate diversity through the Pleistocene may instead be an artefact of an understudied fossil record. ...
There are more species of lizards and snakes (squamates) alive today than
any other order of land vertebrates, yet their fossil record has been poorly
documented compared with other groups. Here, we describe a gigantic
Pleistocene skink from Australia based on extensive material that includes
much of the skull and postcranial skeleton, and spans ontogenetic stages
from neonate to adult. Tiliqua frangens substantially expands the known
ecomorphological diversity of squamates. At approximately 2.4 kg, it was
more than double the mass of any living skink, with an exceptionally
broad, deep skull, squat limbs and heavy, ornamented body armour.
It probably filled the armoured herbivore niche that land tortoises (testudinids),
absent from Australia, occupy on other continents. Tiliqua frangens and
other giant Plio-Pleistocene skinks suggest that small-bodied groups that
dominate vertebrate biodiversity might have lost their largest and often
most morphologically extreme representatives in the Late Pleistocene,
expanding the scope of these extinctions.
... Among Madtsoiidae, small Late Cretaceous and Paleogene taxa such as Adinophis, Alamitophis, Australophis, Patagoniophis, Herensugea, Nanowana, and Nidophis (Albino, 1986(Albino, , 1994(Albino, , 2000(Albino, , 2007Rage, 1996;Scanlon, 1997Scanlon, , 2005Vasile et al., 2013;Pritchard et al., 2014), are markedly smaller, with an estimated body length of less than 2 meters, and most of them have more elongate vertebrae. In contrast, the wide and short, large-sized vertebrae of Powellophis resembles medium-sized taxa such as Rionegrophis, Madtsoia camposi, Menarana nosymena, and Sanajeh (Albino, 1986(Albino, , 2007Rage, 1998;LaDuke et al., 2010;Wilson et al., 2010), which are estimated between 2 and 4 meters in length, and large-sized to gigantic taxa that exceed 4 meters in length such as Gigantophis, Wonambi naracoortensis, Menarana laurasiae, Yurlunggur, Eomadtsoia, Platyspondylophis, and remaining species of Madtsoia (Simpson, 1933;Smith, 1976;Scanlon, 1992;Rage, 1996;Scanlon and Lee, 2000;LaDuke et al., 2010;Mohabey et al., 2011;Rio and Mannion, 2017;Smith et al., 2016;Gómez et al., 2019). The narrow and well-demarcated hemal keel of Powellophis and the lack of diapophyses extending laterally beyond the limit of the prezygapophyses, clearly distinguishes it from those taxa, except the older Eomadtsoia (Late Cretaceous of Patagonia). ...
The Madtsoiidae are an extinct lineage of snakes known from the Late Cretaceous to the Late Pleistocene, with a rich fossil record distributed mainly across Gondwanan landmasses. However, only a few taxa are represented by cranial or articulated remains, and most madtsoiids are known only by isolated vertebrae. The unambiguous record of Madtsoiidae from the Cenozoic in South America had been restricted to the genus Madtsoia from Eocene and Oligocene deposits of Patagonia and Brazil. Here, we describe a new madtsoiid taxon, Powellophis andina gen. et sp. nov., based on an articulated postcranial skeleton from the Mealla Formation (middle–late Paleocene) in northwestern Argentina. The new taxon is estimated to be around 3 meters long, with a vertebral morphology sharing similar features with other mid-to-large forms. Its inclusion in a recent analysis of madtsoiid relationships recovers Powellophis as an early member of a clade formed by mostly large bodied and gigantic taxa. Its presence in the Paleocene of northwestern Argentina fills the gap between the diverse Late Cretaceous and Eocene–Oligocene records of madtsoiids in South America, confirms their presence in northern Gondwana by the early Paleogene, and expands the diversity of the group.
... Jaw bones of agamids are commonly recovered from fossil deposits, but their identification beyond family level is difficult. Some studies state the resemblance of a fossil compared with extant species, without details about characters or criteria used to achieve affiliations (Smith, 1976;Archer et al., 2006;. The relative prevalence of agamid jaw bones in deposits compared to other bones has led to some comparative work on these elements. ...
... Scanlon et al. (2003) cited multiple qualitative characters of the maxilla that unite Incongruelaps with oxyuranines (sensu and other elapids, and the aforementioned characters unite the taxon with Oxyuraninae to the exclusion of Laticauda. The Australian fossil record of Elapids extends from the Quaternary through the Neogene (e.g., Smith, 1976;Scanlon, 2003). The oldest record was previously considered to be latest Paleogene based on a precloacal vertebra from the RSO site of Riversleigh (Scanlon et al., 2003); however, RSO has been radiometrically dated to 16.55 ± 0.29 Ma (Woodhead et al., 2016). ...
... Another giant snake, nearly as large as Yurlunggur, was present in southern Australia well within the last million years; it was the first extinct snake described in Australia, Wonambi naracoortensis (Smith, 1976). More complete material of this species has been found since (Barrie, 1990;Scanlon, 1995b;Scanlon and Lee, 2000). ...
... The other giant Australian madtsoiid was the first extinct snake described in Australia, from the Pleistocene deposits of Naracoorte Caves, South Australia (Smith 1976). Additional remains of Wonambi naracoortensis have been found at Naracoorte and in a number of other sites, all of them limestone cave deposits in the south of the continent (Fig. 3.6; Scanlon 2005a). ...
Large carnivorous reptiles have been an important part of non-marine ecosystems in Australia throughout the Cenozoic, as far as represented in the fossil record. For most groups, this is no earlier than the late Oligocene (roughly 25 million years ago). Fossil deposits of this age contain the earliest known examples in Australia of terrestrial mekosuchine crocodiles, large madtsoiid snakes, pythons, and varanid lizards, alongside the oldest known members of important carnivorous marsupial lineages (thylacines, thylacoleonids, and propleopine kangaroos); crocodyline crocodiles and elapid snakes arrive later. This chapter briefly reviews the fossil history and palaeobiology of each of these carnivorous reptilian groups, with an emphasis on large (megafaunal) species.
... Relatively few sites have yielded squamate remains, and these are generally fragmentary and difficult to interpret given the lack of detailed data on the osteological characteristics of the extant fauna. The best record is that of the Plio-Pleistocene, with cave deposits at several sites yielding fossil lizard and snake remains, which have been studied to varying degrees, although yet to be fully descibed for any location (e.g., Smith 1976, Mackness & Hutchinson 2000, Hutchinson & Mackness 2002, Hocknull et al. 2007, Prideaux et al. 2007, Hollenshead et al. 2010. Most fossil squamate specimens from these sites can be allocated to living genera and species groups, with only a few extinct taxa described so far, e.g., Megalania (now in Varanus;after Hecht 1975), Wonambi (Scanlon 1995) and Aethesia (Hutchinson & Scanlon 2009). ...
Čerňanský, A. & Hutchinson, M.N., 2012. A new large fossil species of Tiliqua (Squamata; Scincidae) from the Pliocene of the Wellington Caves (New South Wales, Australia). Alcheringa, 1–6. ISSN 0311-5518.We describe an isolated frontal bone referable to a new species, Tiliqua laticephala (Scincidae), from the Pliocene Big Sink doline of the Wellington Caves, central eastern New South Wales, Australia. The bone is very robust, is unusually broad and thick, especially around the bases of the subolfactory processes and represents a large and heavily built lizard. The fossil has multiple fragmentary osteoderms in the frontal region, showing asymmetry in shape and thickness that do not correspond to the more regularly arranged anterior head shields of other scincoids. The specimen shares two unusual character states with extant Tiliqua, especially the large armoured species, T. rugosa. Other large skinks related to T. rugosa (other Tiliqua spp., Corucia zebrata, Egernia cunninghamii, Bellatorias major, Liopholis kintorei) are less similar in terms of frontal shape, thickness, sculpture, osteoderm ornamentation, and positioning of adjacent bones.
... In spite of its fragmentary preservation, MPEF-PV 1464 is characterized by this state. The large (centrum length > 18 mm) excludes it from affinity with the smaller-bodied madtsoiid genera Alamitophis Albino, 1986, Patagoniophis Albino, 1986, Rionegrophis Albino, 1986, Herensugea Rage, 1996, Nanowana Scanlon, 1997, Menarana LaDuke et al., 2010and Nidophis Vasile et al., 2013 Smith, 1976(Simpson 1933, Hoffstetter 1961, LaDuke et al. 2010. These latter genera have a series of small parazygantral foramina lateral to the zygantrum, whereas Madtsoia usually have a single, large foramen recessed in a distinct fossa (LaDuke et al. 2010). ...
Squamata are known from South America since the Cretaceous, but their fossil record has an occurrence gap between the late Eocene and early Miocene. Fossils recovered from the Sarmiento Formation (Deseadan South American Land Mammal Age, late Oligocene) at Cabeza Blanca (45°S) partially fill this interval. The squamates recovered from Cabeza Blanca include both lizards (an indeterminate Iguanidae and a probable Iguaninae) and snakes (Madtsoiidae). If these taxonomic assignments are correct, the presence of an Iguaninae at such a latitude is unexpected because these lizards are presently absent from Argentine territory. The madtsoiid, here referred to Madtsoia, would extend the Cenozoic record of this genus back
to around 16 Ma. The squamate fauna from Cabeza Blanca is compatible with warm and humid environments inferred for the Patagonian Deseadan.
... Until now, the fossil history of elapids in Australia has been extremely limited, very recent, and of little practical relevance to questions of the age and pattern of the radiation. Pleistocene elapid fossils have been described from Victoria Cave at Naracoorte, South Australia (referred to the extant genera Pseudonaja, Pseudechis and Notechis; Smith, 1975Smith, , 1976Reed and Bourne, 2000) and reported from the fluviatile Wyandotte Formation, North Queensland (not identified beyond "probably elapid"; McNamara, 1990). Further elapid remains are known from Pleistocene deposits in Queensland (Darling Downs, Floraville, and Riversleigh), New South Wales (Wellington Caves), Victoria (Bacchus Marsh), and cave sites in the southwest of Western Australia (Scanlon, 1995 and unpublished data). ...
Vertebral and cranial remains of elapid snakes have been collected from fossil assemblages at Riversleigh, north-west Queensland, Australia; most are Miocene but one may be late Oligocene and another as young as Pliocene. The oldest specimen (probably the oldest elapid yet known anywhere) is a vertebra that can be referred provisionally to the extant taxon Laticauda (Hydrophiinae, sensu Slowinski and Keogh, 2000), implying that the basal divergences among Australasian hydrophiine lineages had occurred by the early Miocene, in contrast to most previous estimates for the age of this geographically isolated adaptive radiation. Associated vertebrae and jaw elements from a Late Miocene deposit are described as Incongruelaps iteratus nov. gen. et sp., which has a unique combination of unusual derived characters otherwise found separately in several extant hydrophiine taxa that are only distantly related. Associated vertebrae from other sites, and two parietals from a possibly Pliocene deposit, suggest the presence of several other taxa distinct from extant forms, but the amount of material (and knowledge of variation in extant taxa) is currently insufficient to diagnose these forms. The Tertiary elapids of Riversleigh thus appear to be relatively diverse taxonomically, but low in abundance and, with one exception, not referable to extant taxa below the level of Hydrophiinae. This implies that the present diversity of hydrophiine elapids (31 recognized terrestrial genera, and approximately 16 marine) represents the result of substantial extinction as well as the “cone of increasing diversity” that could be inferred from phylogenetic studies on extant forms.
... Skinks of Miocene and Pliocene age have been described from the eastern side of the continent (see discussions and references by Hutchinson 1992, Shea & Hutchinson 1992, Mackness & Hutchinson 2000, Hutchinson & Mackness 2002, Martin et al. 2004, Hocknull 2005, Hutchinson & Scanlon 2009). Late Pleistocene and Holocene sites containing lizard faunas are numerous in South Australia (Tindale 1933, Mulvaney et al. 1964, Thorne 1971, Smith 1976, 1982a,b, Williams 1980, Pledge 1990, Brown & Wells 2000, Reed & Bourne 2000, Queensland (Trezise 1970, Bartholomai 1977, Archer & Brayshaw 1978, Molnar 1978, Horton 1981 and New South Wales (Marshall 1973, Ryder 1974, Dodson et al. 1993, Balme 1995, Dawson & Augee 1997. Surprisingly, Pleistocene lizards from the vast area of Western Australia are largely undescribed. ...
Research on squamate evolution of Australia has predominantly focused on the eastern portion of the continent, whereas little is known about the record from Western Australia. Deposits in Devils Lair provide a glimpse of late Pleistocene Egernia group skink representation from the Cape Leeuwin-Naturaliste region of southwestern Western Australia. Previous studies of fossils from the cave indicate a peak in the number of lizard remains during the last glacial maximum, a time when mammal and charcoal remains suggest an open, dry environment. Previous studies of Devils Lair have not attempted to identify lizard remains beyond ‘lizard’. Here, we identify Egernia group lizard remains as Liopholis spp., Egernia spp., Lissolepis spp. and Tiliqua rugosa, which range in age from ca 48 000 to 13 000 yr B.P.
... Varanid and elapid vertebrae were very common and avian remains were very rare. The extinct giant snake, Wonambi naracoortensis (Smith, 1976), was found only in Layer 3. ...
Grant Hall chamber in Victoria Fossil Cave, Naracoorte, South Australia, contains a late Pleistocene faunal assemblage, dated at between 206 and 76 Ka. Taphonomic and faunal analyses indicate that the predominant mode of accumulation was via a surface exposed pitfall trap. An avian predator, such as Tyto alba, may have been responsible for the accumulation of small mammal remains. The faunal assemblage is taxonomically diverse containing at least 47 taxa. It includes many browsing species such as Wallabia bicolour and the extinct Sthenurine kangaroos and Zygomaturus trilobus, as well as small mammals that require trees and a thick understorey. The Grant Hall fauna thus indicates the presence of densely vegetated woodland, interspersed with small patches of open and thickly grassed areas in the proximal vicinity of the old cave entrance. The relative abundances and species composition of the Macropodidae fauna in Grant Hall are significantly different from other faunal assemblages found at Naracoorte. This study has provided palaeoecological information for a time period not previously investigated at the Naracoorte Caves; detailed surveying of the chamber was undertaken as part of the study.
... Poorly preserved vertebrae were found inside extensions of termite nests. Fossil bone excavated from the Naracoorte cave systems is generally very fragile (Smith, 1971). In the case of the Naracoorte specimens of Wonambi, removal of the matrix from the fossil bones was found to be relatively easy (Barrie, 1990). ...
The skeletal material of Wonambi naracoortensis has been re-examined, and used to test its phylogenetic relationships. The skeletal description, and the resulting data matrix, concentrates on characters that can be assessed by observation (on the associated material of Wonambi kept at the South Australian Museum). Macrostomatan affinities of Wonambi prove to be well supported. The taxon nests within Macrostomata, above Xenopeltis and Loxocemus.
... At least three species have been identified, none of which can be referred to any extant genus (Scanlon 1996). Fossil elapids from Pleistocene deposits in Naracoorte include the modern genera, Pseudechis, Notechis and Pseudonaja (Smith 1975(Smith , 1976). ...
Evolutionary relationships within the Elapidae have been examined using a wide range of morphological and molecular data sets. Historically, there has been little consensus regarding relationships within the largest of the elapid subfamilies, the Hydrophiinae (sensu Slowinski et al. 1997). This presumed monophyletic group includes the sea kraits (Laticauda), the viviparous sea snakes and the terrestrial Australasian elapids. To examine relationships within this subfamily, mitochondrial DNA (partial 12S rRNA) sequence data were obtained for 19 elapid genera including an African, an Asian, 14 terrestrial Australian, a sea krait (Laticauda) and 2 viviparous sea snake genera.
The results are supportive of the following: 1) the subfamily Hydrophiinae (sensu Slowinski et al 1997) is monophyletic, 2) the Australian terrestrial elapids and viviparous sea snakes or ‘palatine draggers’ (McDowell 1970) form a monophyletic group, 3) the sea krait Laticauda is a sister group to the ‘palatine draggers’, 4) there were two separate invasions of the marine environment within the Elapidae, 5) the Australian terrestrial “swamp snake” genus Hemiaspis is a sister group to the viviparous sea snakes indicating that Australia is the centre of origin for this group, 6) the viviparous sea snakes are part of a large, monophyletic clade of viviparous Australian elapids. The viviparous Australian elapids Denisonia devisi and Echiopsis curta may not be part of this radiation, however more work is needed before their relationships within the Hydrophiinae can be resolved. Hypotheses for the origin of Australia’s elapids are considered. The 12S data in conjunction with previous molecular studies and palaeontological evidence are supportive of a marine origin for Australia’s elapid snakes.
... At least three species have been identified, none of which can be referred to any extant genus (Scanlon 1996). Fossil elapids from Pleistocene deposits in Naracoorte include the modern genera, Pseudechis, Notechis and Pseudonaja (Smith 1975(Smith , 1976). ...
Here we report the discovery of fossils representing partial vertebral column of a giant madtsoiid snake from an early Middle Eocene (Lutetian, ~ 47 Ma) lignite-bearing succession in Kutch, western India. The estimated body length of ~ 11–15 m makes this new taxon (Vasuki indicus gen et sp. nov.) the largest known madtsoiid snake, which thrived during a warm geological interval with average temperatures estimated at ~ 28 °C. Phylogenetically, Vasuki forms a distinct clade with the Indian Late Cretaceous taxon Madtsoia pisdurensis and the North African Late Eocene Gigantophis garstini. Biogeographic considerations, seen in conjunction with its inter-relationship with other Indian and North African madtsoiids, suggest that Vasuki represents a relic lineage that originated in India. Subsequent India-Asia collision at ~ 50 Ma led to intercontinental dispersal of this lineage from the subcontinent into North Africa through southern Eurasia.
We here present a thorough documentation of the vertebral morphology and intracolumnar variation across non-caenophidian
snakes. Our studied sample of multiple individuals covers a large number of genera (67) and species (120), pertaining to almost all
extant non-caenophidian families. Detailed figuring of multiple vertebrae across the trunk, cloacal, and caudal series for many different individuals / taxa documents the intracolumnar, intraspecific, and interspecific variation. An emphasis is given in the trunk-tocaudal transition and the pattern of the subcentral structures in that region of the column. Extant non-caenophidian snakes show an astonishing diversity of vertebral morphologies. Diagnostic vertebral features for extant families and many genera are given, though admittedly vertebral distinction among genera in certain groups remains a difficult task. A massive compilation of vertebral counts for 270 species, pertaining to 78 different genera (i.e., almost all known valid genera) and encompassing all extant non-caenophidian families, is provided based on our observations as well as an extensive literature overview. More particularly, for many taxa, detailed vertebral counts are explicitly given for the trunk, cloacal, and caudal portions of the column. Extant non-caenophidian snakes witness an extremely wide range of counts of vertebrae, ranging from 115 up to 546. A discussion on the diagnostic taxonomic utility and potential phylogenetic value of certain vertebral structures is provided. Comparisons of the subcentral structures of the cloacal and caudal vertebral series are also made with caenophidian lineages. We anticipate that this illustrative guide will set the stage for more vertebral descriptions in herpetological works but will also be of significant aid for taxonomic identifications in ophidian palaeontology and archaeozoology.
We here report on the first madtsoiid snake from the late Oligocene of India (the molasse deposits of Ladakh Himalaya). Madtsoiidae is an extinct group of medium sized to gigantic snakes, members of which were mostly distributed across Gondwana.
As an island continent restricted to the Southern Hemisphere, Australia's physiographic, climatic, and biotic histories are unique. This article reviews the records of Australian vertebrates in the middle Pleistocene, an interval of oscillating climate and high faunal diversity, most conspicuously among the larger species composing the 'Pleistocene megafauna'. Australia is divided into eight provinces (major regions) based on modern climatic zones, and the distributions of vertebrates across these provinces are examined. Given the more complete nature of the records for Australian mammals (especially marsupials) compared with those of other vertebrate groups, these species are considered in most detail. Several major 'time-averaged' distribution patterns are evident, particularly among the herbivorous marsupials. These patterns are likely to have been determined by climatically and edaphically controlled distribution of major vegetation types.
Madtsoiidae is a speciose family of extinct snakes that achieved a wide Gondwanan and trans-Tethyan distribution by the Late Cretaceous, surviving until the late Pleistocene. Gigantophis garstini, the first and largest described madtsoiid, was recovered from the upper Eocene of Fayum, Egypt. The 20 vertebrae that constitute the syntype have only received brief description, hindering the referral of specimens to this taxon and our understanding of madtsoiid interrelationships in general. A detailed redescription of the syntype material demonstrates the validity of Gigantophis, based on two autapomorphies (including a strongly depressed neural canal in posterior trunk vertebrae) and a unique combination of characters. Referred material from the lower Paleocene of Pakistan differs significantly, and we restrict Gigantophis to the middle–late Eocene of North Africa. Using a model of morphological variation in extant snakes, we estimate that Gigantophis was 6.9 ± 0.3 m long. A phylogenetic analysis using the largest sample of putative madtsoiids (20 operational taxonomic units) and a revised and augmented matrix (148 characters) places Gigantophis as sister taxon to the latest Cretaceous Indian snake Madtsoia pisdurensis. Whereas our topology might suggest that a dispersal route was present between India and North Africa in the latest Cretaceous–early Paleogene, an evaluation of putative dispersal routes leads us to conclude that the paleobiogeography of Madtsoiidae is best explained by a poorly sampled, earlier widespread distribution in Africa, Indo-Madagascar, and South America. In contrast, latest Cretaceous madtsoiid occurrences in Europe might be explicable by trans-Tethyan dispersal from Africa across the Apulian Route.
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Citation for this article: Rio, J. P., and P. D. Mannion. 2017. The osteology of the giant snake Gigantophis garstini from the upper Eocene of North Africa and its bearing on the phylogenetic relationships and biogeography of Madtsoiidae. Journal of Vertebrate Paleontology. DOI: 10.1080/02724634.2017.1347179.
As an island continent restricted to the Southern Hemisphere, Australia's physiographic, climatic, and biotic histories are unique. This article reviews the records of Australian vertebrates in the middle Pleistocene, an interval of oscillating climate and high faunal diversity, most conspicuously among the larger species composing the ‘Pleistocene megafauna’. Australia is divided into eight provinces (major regions) based on modern climatic zones, and the distributions of vertebrates across these provinces are examined. Given the more complete nature of the records for Australian mammals (especially marsupials) compared with those of other vertebrate groups, these species are considered in most detail. Several major ‘time-averaged’ distribution patterns are evident, particularly among the herbivorous marsupials. These patterns are likely to have been determined by climatically and edaphically controlled distribution of major vegetation types.
New late Cainozoic faunal assemblages are preliminarily identified and described from central eastern Queensland. Biocorrelation of the sites has determined that the oldest faunal assemblages are Early Pliocene in age, with younger faunas from the Plio-Pleistocene, late Pleistocene and Holocene. Pliocene faunal assemblages are characterised by rainforest-specialist frog, squamate and mammalian taxa. These include new Pliocene records for frogs; Kyarranus, Lechriodus, Nyctimystes and microhylids, squamates; Cyclodomorphus gerrardii, a new species of Tiliqua and typholopids, and mammals; Bohra sp., Pseudochirulus spp., new petaurids and dasyurids, Dactylopsila, petauroid incertae sedis, Acrobates, Cercartetus, Uromys/Melomys, Mesembriomys and Pogonomys. Ecological signals derived from the faunal assemblages correlate well with dated palynological records from central eastern and northern Queensland (ODP815, Aquarius Well and Lynch's Crater). Combined Early Pliocene palynological and faunal records strongly indicates a nonseasonal, mesothermal, angiosperm-dominant rainforest with emergent gymnosperms at Mount Etna. A Plio-Pleistocene seasonal, open ecology indicated by the palynological record is corroborated by fauna from similar-aged sites, although several rainforest taxa persist. Increasing aridity during the late Pleistocene is suggested by a distinctly arid-adapted faunal assemblage in late Pleistocene sites, including eastern-most records of Tympanocryptis, Macrotis lagotis, Chaeropus ecaudatus, Perameles bougainville, Sminthopsis macroura and Notomys. Faunal succession from the Early Pliocene to Holocene is characterised by the extinction of most rainforest groups by the late Pleistocene, being replaced by more xeric-adapted forms. Several of the Early Pliocene taxa show resilience to extinction by remaining, albeit rare, in the late Pleistocene fauna, probably in local refugia. These include Dendrolagus sp., a new petauroid, Thylogale, Macroderma gigas, Sarcophilus laniarius and Thylacinus. Presence of rainforest murids in the Early Pliocene of Australia significantly predates previous estimates for their dispersal onto mainland Australia.
The Deccan Traps of peninsular India, representing one of the largest flood basalt eruptions on the earth's surface, have
been a subject of intensive research in the last three decades because of the attributed link between the Deccan Traps and
the Cretaceous-Tertiary boundary mass extinctions. In this context, the biota from the sedimentary beds intercalated with
the volcanic flows and underlying the oldest volcanic flow are more important for understanding the faunal diversity and palaeobiogeography
of India during the time span of volcanic eruptions. A detailed review of the vertebrate faunal diversity of the Deccan volcanic
province is presented here.
Small madtsoiid snakes referred to Patagoniophis and Alamitophis are represented by specimens of all major regions of the vertebral column from the Tingamarra Local Fauna (Early Eocene; Murgon, Queensland, Australia), and are diagnosed as distinct from members of the same genera from the Late Cretaceous (Campanian -?Maastrichtian) of Patagonia, Argentina. The Tingamarra deposit has also produced ribs and dentary bones of two distinct morphotypes, which are provisionally assigned to the same two taxa. A single rib fragment represents a larger form comparable to South American Madtsoia spp., although not positively identifiable as a madtsoiid. It is inferred that two (and probably three) madtsoiid lineages had Antarctic-spanning distributions during the Late Cretaceous and/or Paleocene, with implications for their biology and that of their Neogene and Quaternary descendants.
A new genus and species of extinct lizard is described from the Early Pliocene to Early Pleistocene deposits of Wellington Caves, New South Wales, Australia. The description is based on the anterior half of a left mandibular ramus that shows a suite of unusual characteristics. The fossil is referred to the Scincidae, based on its combination of a distinct coronoid process of the dentary and completely fused Meckelian groove, but it differs significantly from living scincids in several respects. The dental sulcus disappears posterior to the symphysis. The dentition shows slight enlargement of the teeth posteriorly, with the terminal tooth the largest. The new taxon has an exceptionally massive jaw, with a hypertrophied symphysis and coronoid region and the angle at the symphysis suggests a short, deep snout. The new species is not obviously related to any of the major extant scincid lineages.
Here we report for the first time the presence of a pelvic girdle in the fossil snake Wonambi naracoortensis. The material that we attribute to Wonambi consists of a right pelvic girdle, the ilium, pubis, and ischium of which are coossified. It comes from Victoria Cave, a World Heritage Site in the Naracoorte region (about 300 km southeast of Adelaide), South Australia, a locality famous for its richness and diversity of Pleistocene animals...
The lack of an acetabulum at the junction between ilium, pubis, and ischium, together with the lack of an obturator foramen in the pubis, clearly indicates that the hind limbs were either absent or strongly reduced...
Recent expeditions to the Lake Palankarinna area of South Australia resulted in the oldest and only known extinct skink genus in Australia. The holotype of a new genus and species, Proegernia palankarinnensis, was collected from the basal portion of the Late Oligocene Etadunna Formation from the Minkina Local Fauna. Additional scincid fossils previously recovered from higher levels in the formation include material that may be referable to Proegernia. Proegernia is placed in the Egernia group within the Australian Lygosominae, based mainly on its closed Meckelian groove with the apex of the splenial notch low on the lingual surface. However, the apex is markedly more anterior in Proegernia than in any living Egernia group taxon. Proegernia possesses characters suggestive of its position as a stem taxon for later occurring skinks of the Egernia species groups. Even so, overall stage of evolution suggests that the Scincidae of Australia had a long evolutionary history prior to the Late Oligocene, a contention supported by previous molecular studies.
Liasis dubudingala n. sp., described on the basis of isolated vertebrae from the Early Pliocene Bluff Downs Local Fauna, is the largest snake known from Australia. Dependance of vertebral proportions on intracolumnar position indicates that the fossil taxon can be excluded from the Morelia/Python clade. High neural spines suggest possible affinity with Liasis olivacea, whereas a posterior dentary fragment with small teeth is unlike L. olivacea and more similar to Liasis mackloti or species of Bothrochilus and Leiopython. As these extant species have all recently been treated as members of Liasis, the new species is assigned to that genus.
The Central American Loxocemus and the pythons are assigned to sister subfamilies Loxoceminae and Pythoninae, within the weakly characterized family Boidae. They share the character of the left cerebral artery foramen being larger than the right. Within the Henophidia, paired subcaudal scales associate these subfamilies with Xenopeltis and the Uropeltidae.
Twenty-nine discrete variables were coded as 52 binary characters. Nine continuous (meristic) variables were examined. A method is described by which 18 binary characters were selected from seven of these variables, for inclusion in the data set. With Loxocemus and Xenopeltis as outgroups, the data were analysed by a compatibility method. The Pythoninae are resolved into tribes Pythonini and Moreliini on the basis of complementary synapomorphies. The Pythonini have only the genus Python, for the African and Asiatic species, including reticulatus and timoriensis. The Moreliini have all of the Australasian species, in two genera: Aspidites and Morelia.
A phylogenetic analysis of the two tribes is presented, with incomplete resolution of the Moreliini. The two tribes overlap in the areas of the Moluccas and Timor. It is postulated that: Loxocemus plus Pythoninae represent a Laurasian stock; south-east Asia was the primary centre of radiation of the Pythoninae; by the Miocene dispersal to Australasia had occurred, where there was a second radiation.
The limestone caves of the Naracoorte region in South Australia contain extensive deposits of megafauna-rich sediments and intercalated cave formation (speleothems). High-precision thermal ionisation mass spectrometry (TIMS) U/Th dating of flowstones directly associated with several large deposits reveals a distinct cyclicity in the timing of sediment and flowstone depositional events in the Naracoorte caves. This pattern parallels a cyclical alternation of ‘Wet Phases’, and intervening periods with a water deficit over the last 500 ka (Ayliffe et al., 1998. Geology 26, 147). Dates from flowstone interlayers in the fossil deposits coincide with the massive speleothem growth events which characterise Wet Phases. Hiatuses in flowstone deposition correlatable through several cave systems imply that water deficits were initiated by regional aridity during full glacial and then extended into the succeeding warmer, wetter interglacials. Clastic deposits in caverns with restricted entrance shafts correlate with hiatuses, suggesting many of these deposits contain fauna representative of full glacial and interglacial climates. Caverns with small openings have often been sealed from the surface and dating has constrained the length of accumulation episodes in several fossil deposits, one to less than 20 ka. The hydrological regimes and environmental conditions inferred from the timing of speleothem deposition have been used to develop a model for cyclic sediment and bone accumulation in the caverns at Naracoorte over the last 400 ka. The giant Victoria Fossil Chamber deposit accumulated prior to 200 ka and because its entrance is large it may contain faunas representative of all climatic phases. There has been no apparent change in faunal diversity at Naracoorte over at least three glacial–interglacial cycles of the Middle Pleistocene.
A Late Cretaceous (Maastrichtian) assemblage of snakes from the Maevarano Formation of the Mahajanga Basin, northwestern Madagascar, constitutes the only fossil record of snakes from the island. The assemblage, which lived in a highly seasonal, semi-arid climate, includes only archaic forms belonging to the Madtsoiidae and Nigerophiidae, and therefore no representatives of extant Malagasy clades. A large sample of exquisitely preserved vertebrae and several ribs are assigned to Madtsoia madagascariensis, a long (almost 8 m), heavy-bodied ambush predator inferred to have subdued its prey via constriction. A new madtsoiid genus and species, Menarana nosymena, is represented by several associated vertebrae and rib fragments, and part of the basicranium. It was approximately 2.4 m long and appears to have been a powerful, head-first burrower, or at least to have had a burrowing ancestry. Kelyophis hechti, by far the smallest snake in the assemblage (<1 m long), is a new genus and species of primitive nigerophiid based on six isolated vertebral specimens. It was not as specialized for the aquatic lifestyle inferred for other nigerophiids. Although recent molecular phylogeographic studies suggest an early colonization of Madagascar by snakes ancestral to modern Malagasy boids, with subsequent vicariant evolution, the Maevarano Formation assemblage offers no support for this hypothesis. The repeated pattern of extinct archaic lineages being replaced on Madagascar by basal stocks of extant clades (e.g., Anura, Crocodyliformes, Avialae, Mammalia) after the Late Cretaceous is also a plausible scenario for the origin of the extant Malagasy snake fauna.
The notion that Australia's large, terrestrial carnivore faunas of the middle Tertiary to Pleistocene were dominated by reptiles has gained wide acceptance in recent decades. Simple but sweeping hypotheses have been developed seeking to explain this perceived ecological phenomenon. However, a review of the literature does not support these interpretations, which are based on largely speculative and, in many cases, clearly erroneous assumptions. Few size estimates of fossil reptilian taxa are based on quantitative methodology and, regardless of method, most are restricted to maximum dimensions. For species of indeterminate growth, this practice generates misleading perceptions of biological significance. In addition to misconceptions with respect to size, much speculation concerning the lifestyles of large extinct reptiles has been represented as fact. In reality, it has yet to be demonstrated that the majority of fossil reptiles underpinning the story of reptilian domination were actually terrestrial. No postcranial evidence suggests that any Australian mekosuchine crocodylian was less aquatic than extant species, while a semi-aquatic habitus has been posited for madtsoiid snakes and even the giant varanid, Megalania. Taphonomic data equivocally supports the hypothesis that some Australian mekosuchines were better adapted to life on land than are most extant crocodylians, but still semi-aquatic and restricted to the near vicinity of major watercourses. On the other hand, the accelerating pace of discovery of new large mammalian carnivore species has undermined any prima facie case for reptilian supremacy regarding pre-Pleistocene Australia (that is, if species richness is to be used as a gauge of overall impact). However, species abundance and consumption, not richness, are the real measures. On this basis, even in Pleistocene Australia, where species richness of large mammalian carnivores was relatively low, available data expose the uncommon and geographically restricted large contemporaneous reptiles as bit players. In short, the parable of a continent subject to a Mesozoic rerun, wherein diminutive mammals trembled under the footfalls of a menagerie of gigantic ectotherms, appears to be a castle in the air. However, there may be substance to some assertions. Traditionally, erratic climate and soil-nutrient deficiency have been invoked to explain the perception of low numbers or relatively small sizes of fossil mammalian carnivore taxa in Australia. But these arguments assume a simple and positive relationship between productivity, species richness and maximum body mass and either fail to recognise, or inappropriately exclude, other factors. Productivity has undoubtedly played a role, but mono-factorial paradigms cannot account for varying species richness and body mass among Australia's fossil faunas. Nor can they explain differences between Australian fossil faunas and those of other landmasses. Other factors that have contributed include sampling bias, a lack of internal geographic barriers, competition with large terrestrial birds and aspects of island biogeography unique to Australia, such as landmass area and isolation, both temporal and geographic. ZO010 53 S. W ro e Eco lo gy of Au s tr al ia n fos s il ca rn ivor es
A diverse Pleistocene fossil assemblage was recovered from a site (QML1396) exposed in the southern banks of Kings Creek, Darling Downs, southeastern Queensland. The site includes both high-energy lateral channel deposits and low-energy vertical accretion deposits. The basal fossil-bearing unit is laterally extensive, fines upward and its geometry and sedimentary structures suggest deposition within a main channel. The coarse channel fill passes upward into overbank levee deposits made up of lenticular sandy-shelly strata alternating with muds. Several taphonomic biases relating to preservation of different faunal groups and skeletal elements was discerned. Biases may be related to fluvial sorting of the assemblage, but causes for differences between the preservation and accumulation of mammal versus non-mammal terrestrial vertebrates remain unclear. In general, the vertebrate material was accumulated and transported into the deposit from the surrounding proximal floodplain. The assemblage is composed of 44 species including molluscs, teleosts, anurans, chelids, squamates, and small and large-sized mammals. Palaeoenvironmental analysis suggests that a mosaic of habitats, including vine thickets, scrublands, open sclerophyllous woodlands interspersed with sparse grassy understories, and open grasslands, were present on the floodplain during the late Pleistocene. From sedimentological and ecological data, it is evident that increasing aridity during the late Pleistocene led to woodland and vine thicket habitat contraction, and grassland expansion on the floodplain. At present, there is no evidence to support the suggestion that the retraction of late Pleistocene Darling Downs habitats was due to anthropogenic factors.
The Madtsoiidae were medium sized to gigantic snakes with a fossil record extending from the mid-Cretaceous to the Pleistocene, and spanning Europe, Africa, Madagascar, South America and Australia. This widely distributed group survived for about 90 million years (70% of known ophidian history), and potentially provides important insights into the origin and early evolution of snakes. However, madtsoiids are known mostly from their vertebrae, and their skull morphology and phylogenetic affinities have been enigmatic. Here we report new Australian material of Wonambi, one of the last-surviving madtsoiids, that allows the first detailed assessment of madtsoiid cranial anatomy and relationships. Despite its recent age, which could have overlapped with human history in Australia, Wonambi is one of the most primitive snakes known--as basal as the Cretaceous forms Pachyrhachis and Dinilysia. None of these three primitive snake lineages shows features associated with burrowing, nor do any of the nearest lizard relatives of snakes (varanoids). These phylogenetic conclusions contradict the widely held 'subterranean' theory of snake origins, and instead imply that burrowing snakes (scolecophidians and anilioids) acquired their fossorial adaptations after the evolution of the snake body form and jaw apparatus in a large aquatic or (surface-active) terrestrial ancestor.
Relationships between the major lineages of snakes are assessed based on a phylogenetic analysis of the most extensive phenotypic data set to date (212 osteological, 48 soft anatomical, and three ecological characters). The marine, limbed Cretaceous snakes Pachyrhachis and Haasiophis emerge as the most primitive snakes: characters proposed to unite them with advanced snakes (macrostomatans) are based on unlikely interpretations of contentious elements or are highly variable within snakes. Other basal snakes include madtsoiids and Dinilysia--both large, presumably non-burrowing forms. The inferred relationships within extant snakes are broadly similar to currently accepted views, with scolecophidians (blindsnakes) being the most basal living forms, followed by anilioids (pipesnakes), booids and booid-like groups, acrochordids (filesnakes), and finally colubroids. Important new conclusions include strong support for the monophyly of large constricting snakes (erycines, boines. pythonines), and moderate support for the non-monophyly of the trophidophiids' (dwarf boas). These phylogenetic results are obtained whether varanoid lizards, or amphisbaenians and dibamids, are assumed to be the nearest relatives (outgroups) of snakes, and whether multistate characters are treated as ordered or unordered. Identification of large marine forms, and large surface-active terrestrial forms, as the most primitive snakes contradicts with the widespread view that snakes arose via minute, burrowing ancestors. Furthermore, these basal fossil snakes all have long flexible jaw elements adapted for ingesting large prey ('macrostomy'), suggesting that large gape was primitive for snakes and secondarily reduced in the most basal living foms (scolecophidians and anilioids) in connection with burrowing. This challenges the widespread view that snake evolution has involved progressive, directional elaboration of the jaw apparatus to feed on larger prey.
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