Battenizyga, a new Early Triassic gastropod genus with a discussion of the caenogastropod evolution at the Permian/Triassic boundary

Battenizyga, a new Early Triassic gastropod genus from the Moenkopi Formation of Utah, is described and the speciesAnoptychia eotriassica Batten & Stokes, 1986 is placed in it. The new genus has an axially ribbed planktonic larval shell and a teleoconch with an angulated periphery. This character combination is unknown from the Palaeozoic. Therefore,Battenizyga represents additional evidence that recovery from the end-Permian mass extinction was connected with a faunal turnover. Additionally, the extinction of diverse Palaeozoic groups of the Caenogastropoda in the Permian (e.g., the Pseudozygopleuridae) suggest a turnover. All caenogastropod genera that hold Early Triassic species, have post-Palaeozoic type species and most were not reported from the Palaeozoic. This corroborates the view that there was an intense faunal turnover within the Caenogastropoda.Battenizyga is probably a caenogastropod that is closely related to the superfamily Zygopleuroidea which is abundant in the late Palaeozoic and early Mesozoic.      

Smithian (Early Triassic) ammonoids from Tulong, South Tibet

Intensive sampling of the Tulong Formation in South Tibet has facilitated the construction of a highly-resolved middle and late Smithian ammonoid succession. The new biostratigraphical sequence comprises the middle Smithian Brayardites compressus beds, Nammalites pilatoides beds, and the Nyalamites angustecostatus beds followed by the late Smithian Wasatchites distractus beds and Glyptophiceras sinuatum beds. This faunal succession correlates very well with that of other Tethyan sequences such as the Salt Range (Pakistan), Spiti (India), Oman and South China. The Smithian faunal sequence from Tulong contains several taxa with broad geographic distribution (e.g., Owenites, Paranannites spathi, Shigetaceras, Wasatchites), thus enabling correlation with faunal successions from areas outside the Tethys (e.g., USA, British Columbia, Arctic Canada, South Primorye, Siberia). Early Smithian ammonoid faunas are almost absent in Tulong because of a preservation bias (absence of carbonate rocks). Five new ammonoid genera (Brayardites, Nammalites, Nyalamites, Shigetaceras, Tulongites) and six new species (Brayardites crassus, Brayardites compressus, Prionites involutus, ?Subflemingites compressus, Tulongites xiaoqiaoi, Urdyceras tulongensis) are described.     

The biogeography of Early Triassic ammonoid faunas: Clusters, gradients, and networks

The aim of this paper is to quantitatively investigate the spatial and temporal biogeographical relationships of the recovery of ammonoid faunas after the Permian-Triassic mass extinction using three complementary numerical approaches among which is a new, non-hierarchical clustering strategy. The faunal data set consists of a taxonomically homogenised compilation of the spatial and temporal occurrences of ammonoid genera within 20 Early Triassic Tethyan and Panthalassic sites ranging from 40°S to 70°N in palaeolatitudes. In addition to hierarchical cluster analysis (hCA) and nonmetric multidimensional scaling (NMDS), we introduce a third, new non-hierarchical clustering technique allowing the visualisation of a nonmetric interassemblages similarity structure as a connected network constructed without inferring additional internal nodes. The resulting network, which we call a “Bootstrapped Spanning Network” (BSN), allows the simultaneous identification of partially or totally nested as well as gradational linear or reticulated biogeographical structures.The identified interlocalities relationships indicate that the very beginning of the Early Triassic (Griesbachian) corresponds to a very simple biogeographical context, representing a time of great cosmopolitanism for ammonoids. This context shifts rapidly to a more complex configuration indicative of a more endemic and latitudinally-restricted distribution of the ammonoids during the middle and late Early Triassic (Smithian and Spathian). From an evolutionary dynamic point of view, our results illustrate a very rapid (less than ca. 1.4 myr) Early Triassic recovery of the ammonoid faunas, in contrast to many other marine organisms. This recovery is linked with a marked increase in the overall biogeographical heterogeneity, and parallels the formation of a latitudinal gradient of taxonomic richness, which may be essentially controlled by the progressive intensification of the gradient of sea surface temperature. From a methodological point of view, we show that a BSN is a simple, intuitively legible picture of the nested as well as gradational taxonomic similarity relationships, hence providing a good synthesis (and additional insights) between hierarchical clustering and ordination in reduced space results.     

Early Triassic bivalves from the Feixianguan Formation in Xingyi,Guizhou and the Ximatang Formation in Qiubei,Yunnan (southern China)

Based on newly collected materials from the Lower Triassic Feixianguan Formation of Xiongwu section in Xingyi, Guizhou and the Ximatang Formation of Ximatang section in Qiubei, Yunnan, southern China, nine species belonging to seven genera are described. Two bivalve assemblages are recognized and regionally correlated in South China. The bivalve assemblage from the Feixianguan Formation of Xiongwu exhibits a higher diversity including seven species belonging to seven genera: Claraia griesbachi, Leptochondria virgalensis, Entolium (Entolium) microtis, Towapteria scythicum, Bakevillia exporrecta, Bositra sp., and Unionites? fassaensis. In contrast, the bivalve assemblage from the Ximatang Formation of Ximatang has a much lower diversity, consisting of only two species, i.e., Claraia griesbachi and Claraia radialis. Additionally, C. griesbachi and C. radialis, as the most common species of Claraia in the Early Triassic, are revised.     

The First Record of the Pterosaur Ichnogenus Purbeckopus in the Late Berriasian (Early Cretaceous) of Northwest Germany

A hypichnium of a manus imprint (preserved as plaster cast) indicates for the first time the presence of the large pterosaur ichnotaxon Purbeckopus cf. pentadactylus Delair, 1963 in the late Berriasian of northwest Germany. It is only the second record of Purbeckopus globally and the first pterosaur track from Germany. It provides evidence of a very large pterosaur (wingspan c. 6 m) in this area and from this time period not yet represented by skeletal remains. When compared with the English type material, the specimen exhibits some differences that are related mostly to different properties of the substrate on which both were left. These include, in the German track, an impression of the metacarpo-phalangeal joint of the wing finger, normally not present in pterosaur tracks. Also interesting is the rather blunt termination of the deeply impressed digits I–III, indicating rather short and blunt claws, which seem more suitable for walking than for grasping or climbing. The specimens of Purbeckopus in England and Germany occur in different environments: the English locality was situated close to a brackish lagoon, while the German site belongs to a limnic-deltaic system at the margin of a large, freshwater lake.     

Ammonoid age control of the Early Triassic marine reptiles from Chaohu (South China)

Here we described a series of ammonoid specimens from the Early Triassic of Chaohu, South China and recognized the occurrence of Procolumbites for the first time in this area. The Procolumbites layer is about one meter above the first appearance of Chaohusaurus, indicating that the oldest Chaohusaurus is within the Procolumbites Zone of middle Spathian age. This new age constrain is significantly older than the previously suggested Subcolumbites Zone assignment (early late Spathian). To date, Chaohusaurus is the oldest known ichthyopterygian.     

Extremely distal fluvial sandstone within the playa system of Arnstadt Formation (Norian,Late Triassic), Central Germany

The Wachsenburg Sandstone of Thuringia (Central Germany) occurs within playa deposits of the Arnstadt Formation (Late Triassic, Norian) and furnishes an example of ephemeral river metamorphosis under dryland conditions. Characterized by high flow regime features, the sand-dominated lithofacies constitution exhibits sedimentation by channel processes under the influence of recurring flash floods. Bearing signatures of subaerial exposures, the fining-upward lithofacies cycles are bound by low-angle lateral accretion elements suggesting deposition in a meandering stream. Channel migration in response to point bar expansion and active bank erosion, led to the development of four laterally shifting point bar events. Unimodal palaeocurrent patterns with low variance and azimuthal dispersion support the point bar origin of the Wachsenburg Sandstone. With reduced water budget under largely semi-arid climate, the river progressively became smaller, highly sinuous and ultimately abandoned. The resulting point bar succession was finally covered with sheet flow deposits of over bank origin. The sandstone was deposited during a period of low base-level when the playa system temporarily fell dry.     

High-resolution biochronology and diversity dynamics of the Early Triassic ammonoid recovery: The Smithian faunas of the Northern Indian Margin

Based on new collections of abundant and well preserved material from the Salt Range (Pakistan), Spiti (Northern India) and Tulong (South Tibet), several recent studies focused on the taxonomic revision and detailed biostratigraphy of Smithian ammonoids. In this work, biochronological data for these three well-documented basins are analyzed by means of the Unitary Associations method, resulting in a biochronological scheme of unprecedented high-resolution for the Smithian of the Northern Indian Margin (NIM). Data for each basin are first processed separately, thus yielding three local biochronological zonations. Then, the three sequences are processed together as a regional three-section data set for the construction of an inter-basin sequence at the NIM level. The latter zonation comprises 16 Unitary Associations grouped into 13 zones for the entire Smithian. Analysis of ammonoid diversity dynamics based on this new highly resolved time frame highlights (i) a marked diversification during the early Smithian, (ii) a severe extinction during the late Smithian, and (iii) an overall very high turnover throughout the Smithian. At a global spatial scale and stage resolution, the diversity of Smithian ammonoid genera appears surprisingly high, as highlighted by a previous study. It is shown that at a smaller geographic scale and with the most highly resolved time frame, Smithian ammonoids of the NIM reached their explosive diversity peak essentially through extremely high turnover rates rather than through a classic diversification process of high origination rates coupled with low extinction rates. Based on recently published U/Pb ages, regional apparent total rates of origination and extinction of more than 100 species per My can be inferred for the Smithian ammonoids of the NIM.     

Early Miocene Amphibians (Caudata, Salientia) from the Mokrá-Western Quarry (Czech Republic) with comments on the evolution of Early Miocene amphibian assemblages in Central Europe

The Mokrá-Western Quarry exhibits the rare occurrence of Early Miocene (MN 4) vertebrate fauna within the area of the eastern part of Central Europe. In addition to a rich fauna of reptiles and mammals, two fossiliferous karst joints (Mokrá-Western Quarry, 1/2001 Turtle Joint and Mokrá-Western Quarry, 2/2003 Reptile Joint) yielded a rich fauna of amphibians including 13 amphibian taxa: Salamandridae: Mioproteus sp., Chelotriton sp., type I, Chelotriton sp., type II, Triturus aff. roehrsi, Triturus cf. marmoratus, Triturus sp. (T. cristatus species group), Chioglossa meini, Mertensiella mera, Salamandridae gen. and sp. indet.; Pelobatidae: Pelobates sanchizi; Ranidae: Rana sp. (synklepton Rana esculenta); Bufonidae: Bufo sp. The first records of the West European species Triturus cf. marmoratus and Chioglossa meini are reported from the eastern part of Central Europe indicating the wide distribution of those taxa throughout the whole of Europe as early as MN 4. The oldest known record of Pelobates sanchizi documents the Early Miocene presence of representatives closely related to the extinct Late Oligocene representatives of Pelobates. The slow evolution of amphibian species is documented by the presence of Triturus cf. marmoratus and the oldest known occurrence of the extinct salamander Mertensiella mera.     

Facies architecture of the Stuttgart Formation (Schilfsandstein, Upper Triassic), central Germany, and its comparison with modern Ganga system, India

The Stuttgart Formation (Schilfsandstein) is approximately 50 m thick in Thuringia, representing deposition during the “Mid-Carnian Wet Intermezzo”. Stratigraphically it occurs between the Grabfeld and Weser formations, which formed under arid conditions. It comprises NNE-SSW-trending elongate, anastomosing channelized sand-rich bodies with erosional bases (channel belts) that are several kilometres wide and pass laterally into predominantly mudstones deposited in interfluve areas. The source area of these clastics was the uplifted Norwegian Caledonides. Muddy interfluve facies is dominant in exposures in Thuringia, Central Germany.The Lower Stuttgart Formation has an unconformable base that is locally overlain by metre-thick “Basal Beds”. These consist of grey mudstones and thin sandstones deposited under humid conditions in predominantly shallow brackish water environments after a marine ingression via the Eastern Carpathian/Upper Silesian Gate. The following 30-40 m-grey, fine-grained sandstones, siltstones and mudstones were deposited in fluvial environments in channel belts and interfluve areas under humid conditions. These are followed by predominantly reddish mudstones and sandstones of mainly fluvial origin, deposited under somewhat drier conditions with seasonal droughts. The Upper Stuttgart Formation may be more than 16 m thick; it comprises reddish and grey sandstones and mudstones that were mostly deposited in lake delta settings by recurring flash floods. During the deposition of this unit climate was weakly humid with less prominent seasonal draughts.The modern Ganga Plain of India is an analogue for the depositional setting of the Stuttgart Formation. Climatic conditions in Ganga Plain are humid monsoonal with seasonal droughts and roughly comparable with those interpreted for Mid-Carnian times in Germany. The sandy deposits of incised channel belts and channels and muddy deposits of interfluve areas in the Ganga Plain are comparable with the sandstone-dominated channelized facies and mudstone dominated interfluve facies of the Stuttgart Formation, respectively.     

New Genus and Species of Chirotheroid Tracks in the Detfurth-Formation (Middle Bunter,Lower Triassic) of Central Germany

Archosaur tracks are described that have been found in 1999 in a single track bed in the Dethfurth-Formation of the Middle Bunter (Lower Triassic) near Wolfhagen (Central Germany). The taxonomic evaluation of these tracks requires the erection of the new genus and species Protochirotherium wolfhagense ichnogen. n. ichosp. n. Apart from this a rearrangement of groups of chirotheroid tracks is proposed due to the digital proportions.     

A comparison between the Upper Triassic floras of China and the Rhaeto-Liassic floras of Europe and East Greenland

A survey is given of the geological and geographical distribution of the Upper Triassic floras of China, based chiefly on many articles published in Chinese during the last ten years. This is followed by a comparison between the Upper Triassic flora of China and the Rhaeto-Liassic floras of Europe (Sweden, Germany) and East Greenland. It is pointed out that important floral assemblages (for example that of the Liassic Thaumatopteris schenkii Assemblage-zone) occur considerably earlier in China (especially in S.W. China) than in Europe and East Greenland.     

The Lower Triassic Werfen Formation of the Karawanken Mountains (Southern Austria) and its disaster survivor microfossils, with emphasis on Postcladella n. gen. (Foraminifera, Miliolata, Cornuspirida)

A relatively poorly known sedimentary succession (uppermost Bellerophon Formation and overlying Werfen Formation) including the Permian-Triassic Boundary (PTB) is exposed in the South-Alpine part of the Karawanken Mountains of Southern Austria. As in the Dolomites, sedimentation of the Werfen Formation mainly occurred in a subtidal shelf environment between normal and storm wave base. Muddy sedimentation was periodically interrupted by storm events resulting in the deposition of storm layers. Three sections are accurately studied. All contain the same assemblage of Foraminifera and events from the PTB to the late Early Triassic Meandrospira pusilla zone. Between these two great events, only the cornuspirid foraminiferan Postcladella n. gen. is present. Its populations are described and illustrated here under the name Postcladella kahlori n. comb. In the geological record, many taxa such as P. kahlori, i.e., undivided, tubular, initially planispirally evolute coiled, and later uncoiled, appeared successively. A reappraisal is attempted in order to discriminate the different taxa. These forms are often disaster survivors.     

The Ichnofossils of the Triassic Hope Bay Formation,Trinity Peninsula Group,Antarctic Peninsula

The Triassic Hope Bay Formation (Trinity Peninsula Group) includes a diverse ichnocoenosis in the Puerto Moro succession (Hope Bay, Antarctic Peninsula). The Hope Bay Formation is a thick turbidite succession with a minimum vertical exposure of 533 meters along the Hope Bay coast. The rocks are locally affected by contact metamorphism related to later arc magmatism. The ichnofossils are found mainly in thick- and thin-bedded sandstone-mudstone facies composed of a monotonous repetition of sandstone-mudstone cycles. The sandstones are usually medium grained, massive or parallel laminated; the mudstones are massive and rarely laminated. In the fine-grained rocks, mainly the mudstones, there are distinct densities of bioturbation, and at least six patterns were observed. The following ichnogenera were recognized: Arenicolites Salter 1857, Lophoctenium Richter 1850, Taenidium Heer 1877, Palaeophycus Hall 1847, Phycosiphon von Fischer-Ooster 1858 and Rhizocorallium Zenker 1836. All appear to be feeding-traces. The trace fossil assemblages occur mainly in black mudstones rich in organic material that suggest a low oxygen environment. The stratigraphic interval in which they occur is interpreted as progradational supra-fan lobes with channel fill and levee deposits. The thin-bedded turbidite and mudstone lithofacies, where the ichnofossils are abundant, is interpreted as a distal fan turbidite or levee deposit related to a long-term channel fill. This study is the first significant report of trace fossils in the Hope Bay Formation.     

Microfacies and depositional environment of an Upper Triassic intra-platform carbonate basin: the Fatric Unit of the West Carpathians (Slovakia)

Facies associations of the Rhaetian Fatra Formation from the Veká Fatra Mts. (West Carpathians) were deposited in a storm-dominated, shallow, intra-platform basin with dominant carbonate deposition and variable onshore peritidal and subtidal deposits, with 21 microfacies types supported by a cluster analysis. The deposits are formed by bivalves, gastropods, brachiopods, echinoderms, corals, foraminifers and red algae, ooids, intraclasts and peloids. A typical feature is the considerable variation in horizontal direction. The relative abundance and state of preservation of components as well as the fabric and geometric criteria of deposits can be correlated with depth/water energy-related environmental gradients. Four facies associations corresponding to four types of depositional settings were distinguished: a) peritidal, b) shoreface, above fair-weather wave base (FWWB), c) shallow subtidal, above normal storm wave base and d) above maximum storm wave base. The depositional environment can be characterized as a mosaic of low-relief peritidal flats and islands, shoreface banks and bars, and shallow subtidal depressions. The distribution and preservation of components were mainly controlled by the position of base level (FWWB), storm activity and differences in carbonate production between settings. Poorly or moderately diverse level-bottom macrobenthic assemblages are dominated by molluscs and brachiopods. The main site of patch-reef/biostrome carbonate production was located below the fair-weather wave base. Patch-reef/biostrome assemblages are poorly diverse and dominated by the branched scleractinian coral Retiophyllia, forming locally dm-scale autochthonous aggregations or more commonly parautochthonous assemblages with evidence of storm-reworking and substantial bioerosion by microborings and boring bivalves.Facies types and assemblages are comparable in some aspects to those known from the Upper Triassic of the Eastern and Southern Alps (Hochalm member of the Kössen Formation or Calcare di Zu Formation), pointing to similar intra-platform depositional conditions. The absence of large-scale patch-reefs and poor diversity of level-bottom and patch-reef/biostrome assemblages with abundance of eurytopic taxa indicate high-stress/unstable ecological conditions and more restricted position of the Fatric intra-platform setting from the open ocean than the intra-platform habitats in the Eastern or Southern Alps.     

Pangea Megasequences of Tethyan Gondwana-margin reflect global changes of climate and tectonism in Late Palaeozoic and Early Triassic times—A review

The maximum concentration of continental crust at the Pangea stage is characterized by a specific depositional sequence generally referred to as the Pangea Megasequence. Extending in time from the Late Carboniferous to the middle of the Triassic, the succession exhibits similar trends across the whole of Gondwana. Invariably, the sequence was initiated by Late Carboniferous to Early Permian glacial and periglacial deposits. Deglaciation occurred in Early Sakmarian time, evidenced by a typical, commonly transgressive facies. The succeeding formations comprise, in ascending order, coal measures, redbeds, some more coal measures and again redbeds with an intercalation of fluviatile sands in the Early Triassic.After deglaciation the basic depositional theme was modified, depending on postglacial adjustments of climate and on the type of regional tectonic regimes. Extension of the tropical climatic belt after deglaciation was one factor that governed the resulting sediment facies. Coal deposition that prevailed in central Gondwana in the Early Permian gave way to dominance of redbeds in the Middle and Late Permian and, in more distal positions, evaporitic deposits were laid down, following deglaciation. Within marine realms, coralline limestones were formed.Within Gondwana the depositional period of the Pangea Megasequence was governed by three distinctive tectonic regimes: collision dominated the Panthalassa margin, transpressional sag controlled the interior basins, and extension and rifting was experienced along the entire Tethyan margin. In the Early Permian, large and complex graben structures commenced to develop between Africa and India (Malagasy Trough) and between India and Australia (West Australian Trough), giving access to Tethyan waters during deglaciation, commencing in the late Early Sakmarian.Rifting along the Tethyan margin commenced in the Early Permian and was associated with active volcanism between Cashmere and Yunnan and in north-western Australia. Spreading of Neo-Tethys and the formation of oceanic crust, leading to the separation of the Cimmerian Blocks from Gondwana, commenced in the late Early Permian and continued into the Triassic. Thus two facies realms developed, an intracratonic rift facies comprising the Cashmere, Lhasa and Baoshan blocks and a facies controlled by detachment, comprising more distal blocks, such as Tengchong, Malay and Sumatra. The present distribution of individual blocks was governed by fold movements of the Himalayan Orogeny, complicated by transpression along the eastern Himalayan Syntaxis.     

An unusually diverse mollusc fauna from the earliest Triassic of South China and its implications for benthic recovery after the end-Permian biotic crisis

A single carbonate coquinoid lens from the Griesbachian (Early Triassic) of Shanggan, South China, yielded 11 bivalve species described in this study in addition to four gastropod and one ammonoid species reported elsewhere. This makes the Shanggan fauna one of the richest mollusc faunas from the early post-extinction interval after the end-Permian mass extinction event. Four of the present genera are long-term survivors, five are holdovers that went extinct at the end of the Griesbachian or later in the Early Triassic, and seven first appear in the Griesbachian. Three new bivalve species are described: Myalinella newelli nov. sp., Scythentolium scutigerulus nov. sp., and Eumorphotis shajingengi nov. sp. The genus Astartella, previously assumed to have vanished at the end of the Permian, is reported for the first time from the Early Triassic, which also removes Astartidae from Early Triassic Lazarus taxa. The small growth size of the Astartella specimens supports an earlier hypothesis that many of the Early Triassic Lazarus taxa did not survive in unknown refuges but were simply overlooked due to the scarcity of easily observable large-sized specimens. Ecologically, a comparatively high proportion of infaunal bivalve species (4/11) is remarkable for the early post-extinction interval, supporting the impression of a relatively advanced recovery state. Moreover, abundance-data of the bivalve-gastropod community reveal a remarkably low dominance index (D = 0.17) that is suggestive for advanced recovery and stable environmental conditions. It is proposed that the Shanggan fauna represents a late Griesbachian benthic recovery event that coincided with the appearance of similarly diverse benthic faunas in Oman and Primorye. A high proportion of genera that have previously not been reported from the Early Triassic indicate that the prevalence of poor preservation conditions is a major obstacle in identifying early phases of recovery from the greatest crisis in the history of metazoan life. The early recovery of benthic faunas reported in this study questions previous claims of a prolonged lag phase as a consequence of the extraordinary extinction magnitude or the persistence of adverse environmental conditions.     

High-resolution sequence interpretation of epeiric shelf carbonates by means of palynofacies analysis: An example from the Germanic Triassic (Lower Muschelkalk, Anisian) of East Thuringia, Germany

Summary The Steudnitz quarry (East Thuringia, Germany) exposes the entire Lower Muschelkalk (Middle Triassic, Anisian) with a thickness of 103 m. In this paper, a primarily palynofacies-based sequence-stratigraphical interpretation supported by lithological investigations is proposed. Palynofacies of the sampled carbonate rocks is dominated by land-derived phytoclasts and pollen grains. Spores are quite rare. Marine plankton reaches its maximum abundance in an interval around the ThuringianSpiriferina-bed and the Terebratel-beds. In accordance with lithological features, this interval may be interpreted as a third-order maximum-flooding zone. Two different third-order interpretations are presented and discussed. Third-order sequences and parasequence sets can be identified by use of palynofacies analysis. Furthermore, palynofacies analysis provides an additional possibility of recognizing partly eroded or originally incomplete cycles (‘missed beats’). In some cases, a resolution down to the level of parasequences is possible. High-resolution palynofacies analysis is a powerful new tool in sedimentology but should always be combined with lithofacies interpretation. It provides very useful, additional information for high-resolution sequence analysis and palaeoenvironmental interpretation. Being a rather young method, high-resolution palynofacies analysis still shows some calibration problems with classical methods. However, these should be solvable by further research.     

The Triassic reptiles Brachyrhinodon and Polysphenodon and the relationships of the sphenodontids

Two Triassic sphenodontid reptiles, Brachyrhinodon taylori and Polysphenodon mulleri , are redescribed. Both genera are blunt-snouted forms but, as a result of distortion in the original fossils, the degree to which the snouts are shortened has previously been over-elaborated. An elongate temporal region and an interorbital width exceeding that of the parietal table are both derived characters in sphenodontids. Polysphenodon is plesiomorphic for both conditions, whereas Brachyrhinodon displays the derived states. Therefore, it is assumed that the reduced snout has been independently derived in each genus. The limb elements of Polysphenodon are long relative to the size of the skull and are similar in proportions to some species of Homoeosaurus. This resemblance is considered to be indicative of similar locomotory requirements rather than close evolutionary relationships. Cladograms showing possible interrelationships within the Sphenodontida are based on twenty-nine character states.