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Lepidostrobus muelleri Johnston is a skull fragment of a temnospondyl amphibian Andrew C. Rozefeldsa; Anne Warrenb a Tasmanian Museum and Art Gallery, Hobart, Tasmania, Australia b Department of Zoology, La Trobe University, Melbourne, Victoria, Australia

First published on: 01 December 2010

To cite this Article Rozefelds, Andrew C. and Warren, Anne(2010) 'Lepidostrobus muelleri Johnston is a skull fragment of a temnospondyl amphibian', Alcheringa: An Australasian Journal of Palaeontology,, First published on: 01 December 2010 (iFirst) To link to this Article: DOI: 10.1080/03115518.2011.527163 URL: http://dx.doi.org/10.1080/03115518.2011.527163

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The publisher does not give any warranty express or implied or make any representation that the contents will be complete or accurate or up to date. The accuracy of any instructions, formulae and drug doses should be independently verified with primary sources. The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. Palaeontological note p. 185, pl. 22, figs 7–12) noted that the specimen was collected from Mesozoic rocks Lepidostrobus muelleri Johnston is a and was associated with Phyllotheca leaves and stems. It derives from the Upper Par- skull fragment of a temnospondyl meener Supergroup (Farmer 1979) — most amphibian likely from the Lower Triassic Knocklofty Formation or its stratigraphic equivalents. ANDREW C ROZEFELDS AND Johnson interpreted Lepidostrobus muel- ANNE WARREN leri as a reproductive axis with sporangia and narrow chambers. It lacks, however, any evidence of spiral phyllotaxy that would ROZEFELDS, A.C. & WARREN, A., iFirst article. Lepidos- be expected if the specimen were a cone. trobus muelleri Johnston is a skull fragment of a Johnston (1885, 1888) also reported winged temnospondyl amphibian. Alcheringa, 1–4. ISSN 0311- 5518. seeds in the sandstone surrounding the impression but examination, using a com- A fossil described as Lepidostrobus muelleri Johnston is pound microscope, revealed no structures referred to temnospondyl amphibians and is shown to have affinities with either the Rhinesuchidae or the that could be confidently identified as seeds. Rhytidosteidae. It seems likely that Johnston (1885) mis- interpreted the flakes of black mica in the Andrew C Rozefelds ([email protected] sandstone as carbonaceous seeds. v.au), Tasmanian Museum and Art Gallery, GPO Box 1164, Hobart, Tasmania, 7001 Australia; Anne Warren Walkom (1925) was unable to determine ([email protected]), Department of Zoology, La the affinities of this specimen but pointed out Trobe University, Melbourne, Victoria, 3086, Australia. that it was not referable to Lepidostrobus, Received 6.8.2010; revised 6.9.2010; accepted 15.9.2010. mainly because that genus is restricted to Palaeozoic strata. Max Banks, pers. comm., Key words: temnospondyli, Amphibia, Rhinesuchidae, Rhytidosteidae, Triassic, Tasmania. in Crespin (1964), interpreted Z1376 as a probable jaw of an amphibian, although it was not examined or cited in Cosgriff’s (1974) seminal work on the Tasmanian JOHNSTON (1883, 1885) mentioned and temnospondyls. In this note the specimen is then formally described a new species of described, and its affinities are re-examined. Downloaded By: [Rozefelds, Andrew C.] At: 23:37 8 December 2010 Lepidostrobus, a genus used for microspor- A latex cast was made of the impression and angiate cones of Lepidodendron (Lycopsida), examination of the cast confirms Banks’ together with putative scales that were suggestion that Z1376 is an incomplete interpreted as winged seeds. This specimen, double tooth row of a temnospondyl. held in the Tasmanian Museum and Art Gallery (Z1376), is an impression only and was recorded by Johnston (1883) as being Description of skull fragment collected from the thick-bedded sandstones The specimen is from a skull or mandible of Campania in southeastern Tasmania. that was straight-sided rather than rounded The specimen is preserved in a coarse- (Fig. 1A). Approximately 35 tooth loci are grained sandstone and lies on a bedding preserved on the marginal tooth row, plane associated with quartz-pebble and comprising 20 tooth bases and 15 replace- clay-pebble conglomerates. Johnston (1888, ment pits (Fig. 1B). Some teeth show the typical temnospondyl labyrinthine tooth ISSN 0311-5518 (print)/ISSN 1752-0754 (online) Ó 2010 Association of Australasian Palaeontologists structure, consisting of infolded dentine DOI: 10.1080/03115518.2011.527163 layers. The marginal teeth are strongly 2 A. C. ROZEFELDS and A. WARREN ALCHERINGA

Comparisons with temnospondyl amphibians Temnospondyls all have a marginal row of teethonthedentaryofthelowerjawandon the maxilla of the upper jaw. In Triassic temnospondyls (Stereospondyli), lingual to the dentary tooth row on the mandible, are a series of three coronoid bones that in some cases are tooth-bearing and denticulate. All three coronoids bear denticles in the rhine- suchid Rhineceps nyasaensis (Watson 1962), the lydekkerinid Lydekkerina huxleyi (Jeannot et al. 2006, Hewison 2007) and the rhytidosteids (e.g. Arcadia myriadens:War- ren & Black 1985). Except in the Plagiosaur- idae, in which teeth are present in rows on all three coronoids but denticles are absent (Hellrung 2003, Jenkins et al. 2008), coronoid teeth are rarely present on more than the posterior coronoid, and tend to be smaller than the marginal teeth and more scattered. For the above reasons Z1376 is more likely to Fig. 1. Z1376 interpreted as a fragment from the right be from the skull, with a labial maxillary side of a temnospondyl skull. A, photograph of latex cast in ventral view; B, Line drawing of TMAGZ1376. tooth row and lingually the posterior part of D ¼ denticle patch; EC ¼ ectopterygoid; MX ¼ maxilla; a palatine and most of an ectopterygoid. In PAL ¼ palatine; R ¼ replacement pit; T ¼ tooth. Scale Triassic temnospondyls, these inner bones of bar ¼ 1 cm. the palate commonly bear rows of teeth that are of equivalent size to those of the adjacent compressed anteroposteriorly, and their maxilla. Shagreen denticles are uncommon long axes are aligned at an angle to the on the palatine and ectopterygoid, being Downloaded By: [Rozefelds, Andrew C.] At: 23:37 8 December 2010 jaw margin, an unusual orientation (Fig. recorded only in the Rhinesuchidae (Watson 1B). Lingual to the marginal tooth row, two 1962) and Rhytidosteidae (Cosgriff 1964, bones are preserved but separated from one Warren & Black 1985). In those families, another by a distinct angled suture, and well-preserved palates are rare. In the rhine- from the marginal bone by a longitudinal suchid, Rhineceps nyasaensis, the inner teeth suture. The tooth-bearing parts of the of the palate are somewhat irregular in lingual bones are raised above the general arrangement and size on the right, but more surface of the bone (Fig. 1B). These bones regular on the left, and some of the maxillary are more abraded than the marginal bone, teeth are aligned at an angle to the jaw but it is clear that they were tooth-bearing, margin as they are in Z1376 (Watson 1962). the more anterior carrying at least five teeth The arrangement of the palatal bones is with space for eight replacement pits, with a similar to that in Z1376, with the width of the minimal count of four teeth and four pits on combined tooth rows narrow, and an almost the more posterior bone. In addition, a straight margin of the interpterygoid vacuity. shagreen of denticles is present on the Rhinesuchids have not been recorded from anterior third of each bone (Fig. 1B). any Australian deposits. ALCHERINGA TASMANIAN TEMNOSPONDYL 3

Australian members of the Rhytidostei- Tasmania have been little studied and dae have skulls that are either straight- further fieldwork in these areas is warranted. sided, like Deltasaurus kimberleyensis and D. pustulatus (Cosgriff 1964), or parabolic, like Rewana quadricuneata (Howie 1972), Acknowledgements Arcadia myriadens (Warren & Black 1985) The authors thank Steven Forsyth, for and Derwentia warreni (Cosgriff 1974). In advice on the Triassic stratigraphy of Tas- these taxa, the skulls are short with a mania, and acknowledge assistance from the rounded margin of the interpterygoid va- reviewers, Professor Bruce Rubidge and Dr cuity and a proportionally greater width Marcello Ruta, and the Editor, Dr Stephen across the combined marginal and inner McLoughlin, in improving this note. tooth rows than in longer-skulled temnos- pondyls or Z1376. In addition, in these taxa the teeth of the palatine and ectopterygoid References are smaller than the marginal teeth and are COSGRIFF, J.W., 1964. A new genus of Temnospondyli irregular in size. One rhytidosteid with a from the Triassic of Western Australia. Journal straight-sided skull, Pneumatostega potamia of the Royal Society of Western Australia 48, 85– (Cosgriff & Zawiskie 1979) from the Early 90. COSGRIFF, J.W., 1974. Lower Triassic Temnospondyli Triassic of South Africa, is morphologically of Tasmania. Geological Society of America Special closer to the Tasmanian specimen in that Paper 149, 1–134. the combined width of the tooth-bearing COSGRIFF, J.W. & ZAWISKIE, J.M., 1979. A new species bones beside the interpterygoid vacuities is of the Rhytidosteidae from the Lystrosaurus Zone and a review of the Rhytidosteoidea. Palaeontolo- less, and the teeth from the palatine and gia Africana 22, 1–27. ectopterygoid are of a similar size to those CRESPIN, I., 1964. Catalogue of fossil type and figured of the maxilla and are regularly arranged in specimens in Tasmania. Bureau of Mineral a tooth row. Both parabolic (Derwentia Resources, Geology and Geophysics. Report No 69, 30 pp. warreni) and straight-sided (Deltasaurus FARMER, N., 1979. Hobart. Geological Atlas 1:250,000 kimberleyensis) members of the Rhytidos- series, Sheet SK-55/8. Tasmania Department of teidae have been recorded from Tasmania Mines, Geological Survey Explanatory Report, 24 (Cosgriff 1974), but no detail of the palatal pp. HELLRUNG, H., 2003. Gerrothorax pustuloglomeratus, dentition was available in Deltasaurus. Downloaded By: [Rozefelds, Andrew C.] At: 23:37 8 December 2010 ein Temnospondyle (Amphibia) mit kno¨cherner Branchialkammer aus dem Unteren Keuper von Kupferzell (Su¨ddeutschland). Stuttgarter Beitra¨ge Conclusion zur Naturkunde Serie B (Geologie und Pala¨ontolo- gie) 39, 1–130. Z1376 is undoubtedly from a temnospondyl HEWISON, R.H., 2007. The skull and mandible of the amphibian, and could be from a member of stereospondyl Lydekkerina huxleyi (Tetrapoda: the Rhinesuchidae or the Rhytidosteidae. If Temnospondyli) from the Lower Triassic of South Africa, and a reappraisal of the family Lydekker- a rhinesuchid, it is the first record of that inidae, its origin, taxonomic relationships and family from Australia; if a rhytidosteid, it is phylogenetic importance. Journal of Temnospondyl not from a known member of that family. Palaeontology 1, 1–80. While temnospondyls are the most common HOWIE, A.A., 1972. On a Queensland labyrinthodont. In Studies in Vertebrate Evolution, K.A. JOYSEY & vertebrates collected from the Triassic of T.S. KEMP, eds, Oliver & Boyd, Edinburgh, 50– Tasmania, this specimen was found in an 64. area of the State that has yielded undescri- JEANNOT, A.M., DAMIANI,R.&RUBIDGE,B.S.,2006. bed fish bones (Cosgriff 1974) but no other Cranial anatomy of the Early Triassic stereospondyl Lydekkerina huxleyi (Tetrapoda: Temnospondyli) vertebrate fossils. Since the classic research and the taxonomy of South African lydekkerinids. of Cosgriff (1974), the Triassic faunas of Journal of Vertebrate Paleontology 26,822–838. 4 A. C. ROZEFELDS and A. WARREN ALCHERINGA

JENKINS,F.A.JR,SHUBIN, N.H., GATESY,S.M.& JOHNSTON, R.M., 1888. Systematic Account of the WARREN, A., 2008. Gerrothorax pulcherrimus from Geology of Tasmania. William Thomas Strutt, the Upper Triassic Fleming Fjord Formation of Government Printer, Hobart, 408 pp. East Greenland and a reassessment of head lifting WALKOM, A.B., 1925. Notes on some Triassic plants. in temnospondyl feeding. Journal of Vertebrate Papers and Proceedings of the Royal Society of Paleontology 28, 935–950. Tasmania (for the year 1924), 73–89. JOHNSTON, R.M., 1883. Proceedings for June 1882. WARREN,A.&BLACK, T., 1985. A new rhytidosteid Papers and Proceedings of the Royal Society of (Amphibia, Labyrinthodontia) from the Early Tasmania (for the year 1882), p. xxxii. Triassic Arcadia Formation of Queensland, Austra- JOHNSTON, R.M., 1885. Discovery of a cone, probably lia, and the relationships of Triassic temnospondyls. of a species of Lepidostrobus, in the sandstones Journal of Vertebrate Palaeontology 5,303–327. of Campania. Papers and Proceedings of the WATSON, D.M.S., 1962. The evolution of the Labyr- Royal Society of Tasmania (for the year 1884), inthodonts. Philosophical Transactions of the Royal 225. Society of London B 245, 219–265. Downloaded By: [Rozefelds, Andrew C.] At: 23:37 8 December 2010