PROCEEDINGS of NEW SOUTH WALES VOLUME 130 NATURAL HISTORY IN ALL ITS BRANCHES THE LINNEAN SOCIETY OF NEW SOUTH WALES ISSN 0370-047X Founded 1874 Incorporated 1884 The Society exists to promote the cultivation and study of the science of natural history in all its branches. The Society awards research grants each year in the fields of Life Sciences (the Joyce Vickery fund) and Earth Sciences (the Betty Mayne fund), offers annually a Linnean Macleay Fellowship for research, contributes to the stipend of the Linnean Macleay Lecturer in Microbiology at the University of Sydney, and publishes the Proceedings. It holds field excursion and scientific meetings, including the biennial Sir William Macleay Memorial Lecture delivered by a person eminent in some branch of natural science. Membership enquiries should be addressed in the first instance to the Secretary. Candidates for election to the Society must be recommended by two members. The present annual subscription is $456.00. 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Box 82, Kingsford NSW 2032, Australia Telephone: (International) 61 2 9662 6196; (Aust) 02 9662 6196 E-mail: linnsoc @iinet.net.au Home page: http://linneansocietynsw.org.au Cover motif: Reconstruction of Palorchestes from the paper by B.S. Mackness, page 30 this volume. PROCEEDINGS of the LINNEAN SOCIETY of NEW SOUTH WALES For information about the Linnean Society of New South Wales, its publications and activities, see the Society’s homepage http://linneansocietynsw.org.au VOLUME 130 March 2009 u x aE on ihe . ris ry Tali sit i’ AS q 4 : WEAR ORCAS ELT LAT ab Om Aa Wart NO earch cles 4 hey upd rorterrrrd ter 4 SGP Se VIMO Ul Bao earteyeiag ese Arh. Ccmeate CD: ET PTE ert Museum Holdings of the Broad-headed Snake Hoplocephalus bungaroides (Squamata: Elapidae) JAMIE M. HARRIS AND Ross L. GOLDINGAY School of Environmental Science and Management, Southern Cross University, Lismore NSW 2480, Australia; Harris, J.M. and Goldingay, R.L. (2009). Museum holdings of the Broad-headed Snake Hoplocephalus bungaroides (Squamata: Elapidae). Proceedings of the Linnean Society of New South Wales 130, 1-19. The broad-headed snake Hoplocephalus bungaroides (Schlegel, 1837) is a highly endangered species endemic to the Sydney basin. We attempted to track down the whereabouts of museum specimens of this snake by contacting mainly Australian, European and North American curators of natural history museums and university herpetological collections. We received replies from 200 institutions, and from these we present details of 159 specimens from 27 museums in 11 countries reported to us as H. bungaroides. Countries include Australia (108 specimens), Germany (13), the United States (9), United Kingdom (7), France (4), Belgium (5), the Netherlands (5), Austria (3), Denmark (3), Italy (1), and Switzerland (1). At least 47 specimens are from the 19" Century, and accurate locality records were available for 98 specimens. Obviously, all of the specimens have value insofar as they may provide important biological data that will be useful to researchers working on the future conservation of this snake. Many of these specimens also provide important historical evidence of the species’ past distribution. Manuscript received 6 February 2008, accepted for publication 17 September 2008. KEYWORDS: conservation, distribution, museum, reptile, natural history, Krefft INTRODUCTION The broad-headed snake Hoplocephalus bungaroides is possibly the most endangered snake in Australia, with research indicating there are serious concerns for its future conservation (Shine and Fitzgerald 1989; Webb and Shine 1997, 1998a,b; Goldingay 1998; Shine et al. 1998; Goldingay and Newell 2000; Webb et al. 2002; Newell and Goldingay 2005). It has a highly restricted distribution within the Sydney basin where it is dependent on habitats characterised by sandstone cliffs, ridges and outcrops (Krefft 1869; Longmore 1989; Cogger 2000; Swan et al. 2004). This species is threatened by habitat loss through urbanisation, removal of bush rock for landscaping and ongoing degradation of rocky habitat caused by hikers and reptile poachers (Hersey 1980; Shine and Fitzgerald 1989; Cogger et al. 1993; Goldingay and Newell 2000; Webb and Shine 1998a, 2000; Newell and Goldingay 2005). The decline of H. bungaroides was noted as early as 1869 by Gerard Krefft (1830-1881), Curator and Secretary of the Australian Museum (Whitley 1961, 1969), in The Snakes of Australia (Krefft 1869), the first monograph published on Australian snakes. Krefft (1869) considered H. bungaroides (as its junior synonym Hoplocephalus variegatus) to be “very local” with specimens found only “in the immediate neighbourhood of Sydney”, that is, from Port Jackson to Botany Bay, on the shores of Middle Harbour, and at Lane Cove and Parramatta inlets. Krefft stated that this snake is “not so numerous as they were six or eight years ago” (i.e. around 1861-1863) and the decline was attributed to “their haunts having been invaded by the builder and the gardener”. Krefft also stated that “many hundreds” of H. bungaroides specimens had been distributed to unnamed “kindred institutions”. These statements sparked our curiosity, and subsequently we made considerable effort to locate these specimens. In so doing, we also aimed to gather information on all museum holdings of H. bungaroides because this may offer a rich source of data potentially useful to the future conservation of this endangered species. MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE MATERIALS AND METHODS We reviewed the annual reports of the Australian Museum for mention of reptile specimens received and exchanged by Krefft during his tenure as Curator and Secretary (1861-1874) (see Appendix 1). We also searched for information in the archives of the Australian Museum, including examination of Krefft’s correspondence and the ‘Exchange Register’ (pre 1874; series 58, Volume 1). Finally, we surveyed other museums and related institutions with herpetological collections, particularly those in Australia, Belgium, Czech Republic, France, Germany, India, Italy, Netherlands, Portugal, Spain, United Kingdom (UK) and the United States (US) since Krefft did send reptile specimens to these countries (Appendix 1). Museums in these and other countries were identified using online directories and also published lists in Leviton et al. (1980, 1985) and Roselaar (2003). Curators or collection managers were asked via email whether there were any H. bungaroides (or its synonym H. variegates) in their museums. If H. bungaroides was present, data were requested on numbers of specimens held; catalogue / registration numbers; collection locality; collector or donor name; collection date; and other details recorded with the specimens. Additionally, photographs of the specimens were requested to confirm that the correct identifications had been made. In relation to photographs, for one museum in France (Musée de Zoologie, Strasbourg) we received reports about two H. bungaroides specimens in their collection, but the photographs supplied did not reveal the striking appearance of H. bungaroides and we believe they represent the Stephens banded snake H. stephensii. We are confident about the identification of all other museum specimens listed, except for those at Zoological Museum, University of Liege (Belgium) because photographs of the five H. bungaroides in their collection were not supplied. RESULTS The annual reports of the Australian Museum for 1861-1874 did not provide details of “many hundreds” of H. bungaroides. Descriptions of species exchanges in these reports lack detail, and indicate, at a minimum, that H. bungaroides was definitely sent out to only three places (see Appendix 1). The reports mention that reptiles were shipped to a number of museums and specimen dealers in this period, but the specific composition of the shipments was generally not published. Recipients of Krefft’s reptiles included his colleagues in Mauritius (Victor de Robillard) and India (Richard Henry Beddome); one learned society (Royal Society of Tasmania); four specimen ‘dealers’ - J.C. Puls (Belgium), C.L. Salmin (Hamburg, Germany), Vaclav Fri¢ (Prague, Czech Republic), and Robert Damon (Weymouth, England); and at least nine museums, i.e. those in Hamburg and Berlin (Germany), Leiden (Netherlands), Madras (=Chennai, India), Milan (Italy), Paris (France), Madrid (Spain), London (UK), and Harvard at Cambridge (US). The pre-1874 Exchange Register in the Australian Museum archives (series 58, Volume 1) contained some inbound 1860s correspondence addressed to Krefft from dealers such as J.C. Puls and some museums, such as the Muséum National d’Histoire Naturelle, Paris and the Museum of Comparative Zoology, Cambridge. This Register also lists some, but not all, specimens sent on exchange by Krefft and also his predecessor George Bennett. These lists include an entry that a single H. bungaroides was sent to the Government Museum at Madras (=Chennai, India) (see Exchange Register p.16). Whilst this list is undated, it was probably the same consignment listed in the annual report for 1864 (see Appendix 1). The Exchange Register also itemised specimens dispatched to the Royal Society of Tasmania and H. bungaroides was absent from this list. By contacting museums directly, we located 159 specimens reported to us as H. bungaroides from 28 institutions in 11 countries (Table 1). Most specimens we found are held in Australia (108 specimens), but a considerable number are in Europe (43 specimens) and the US (9 specimens). Negative responses to our email enquiries were received from 174 institutions (see Appendix 2). There were also 74 other institutions that did not respond to our correspondence, despite more than one request (Appendix 2). We have compiled some detailed information on H. bungaroides specimens from many institutions in Australia, Europe and the US (see below). Australian collections The Australian Museum, Sydney (AM), has 77 H. bungaroides specimens (Table 1; Appendix 3) but none of these are designated type specimens (Shea and Sadlier 1999). Eight of these do not have any locality data and another 5 have an imprecise collection locality recorded as “Sydney”. There are 18 AM specimens collected at Waterfall, seven at Nowra, five at Long Bay, six at Royal National Park (NP) (including Bundeena), three at the Blue Mountains, three at Woronora Dam and two at La Perouse. Single AM specimen locality records were recorded for 20 locations (Appendix 3). Twenty four (31 %) of the Proc. Linn. Soc. N.S.W., 130, 2009 J.M.HARRIS AND R.L. GOLDINGAY Table 1: Specimens of Hoplocephalus bungaroides held in Australian and overseas museums. Institution Code n Australia Australian Museum, Sydney AM Vi Western Australian Museum, Perth WAM 3 Museum Victoria, Melbourne NMV 6 South Australian Museum, Adelaide SAMA 6 Queensland Museum, Brisbane QM 4 Northern Territory Museum, Darwin NTM 4 Australian National Wildlife Collection, Canberra ANWC 3 Macleay Museum, University of Sydney MMUS 3 Biological Museum, Australian National University ANU 2 Austria Museum of Natural History, Vienna NMW 3 Belgium Zoological Museum, University of Liege MZULG 5 Denmark Zoological Museum, University of Copenhagen ZMUC 3 France Muséum National d’ Histoire Naturelle, Paris MNHNP 4 Germany Museum fiir Naturkunde, Berlin ZMB 8 Senckenberg Natural History Museum, Frankfurt SMF 1 Zoologisches Museum, University of Hamburg ZMH 2 Zoologische Staatssammlung, Munich ZSM 2 Italy Museo Civico di Storia Naturale, Genoa MSNG i ~ Netherlands National Museum of Natural History, Leiden RHNH 5 Switzerland Naturhistorisches Museum, Basel NMB 1 United Kingdom Natural History Museum, London BMNH 6 Oxford University Museum of Natural History OUM 1 United States Field Museum of Natural History, Chicago FMNH 2 Museum of Comparative Zoology, Harvard University MCZ 3 National Museum of Natural History, Smithsonian Institution USNM 2 San Diego Natural History Museum SDNHM 1 University of Illinois Museum of Natural History UIMNH 1 Total 159 Proc. Linn. Soc. N.S.W., 130, 2009 MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE 77 AM specimens do not have collection dates, but presumably some of the undated specimens are very old and derive from the late 19" Century (Krefft’s era). The collection dates on the remaining 53 range from 1904 to 1996. The Macleay Museum (MMUS) holds three H. bungaroides that all are believed to be from the late 19" Century. One is from “Mount Wilson” but the collector and date are unknown. It was possibly John Anderson or James Cox since both of these zoologists made collections for MMUS in the Mount Wilson area (Fletcher 1929; Stuart Norrington pers. comm.). The only information with the two other MMUS specimens is that they were collected on the “coast near Sydney”. Hoplocephalus bungaroides specimens are also held in all other Australian mainland capital cities. The South Australian Museum, Adelaide (SAMA), has 6 specimens recorded on its collection register, but one of these (R00463) is now missing. This misplaced specimen is recorded as collected on 2 June 1915 at La Perouse and donated to the SAMA by the AM. Other SAMA specimens were from Kuringai Chase, Sydney and Woronora River. The Kuringai Chase specimens are reported to us as having been collected by “W. Irvine” in 1967. We enquired with William (Bill) Irvine (a well-known collector who still lives in Sydney) for details about these but he explained that his field notebooks from 40 years ago had now been destroyed. The Queensland Museum, Brisbane (QM), has four specimens: one from Waterfall; one from Nowra that was held in captivity for a period of time (Queensland Reptile Park); one was captive-bred; and another was confiscated by Queensland Parks and Wildlife Service in 1989. In Canberra, three specimens are in the Australian National Wildlife Collection (ANWC): one from about 1963-1964; the other two from around 1978- 1980 (J. Wombey pers. comm.). Collection localities are not available for any of these. Also in Canberra, the Museum at the Australian National University (ANU) has two specimens: Waterfall and Tiajuara Falls (22 km from Nerriga), although these have no dates or registration numbers. In Darwin, the Northern Territory Museum (NTM) has four specimens all from the 1970s and collected at Heathcote, Jarra Fall (Nowra), and Woronora Dam. In Melbourne, Museum Victoria (NMV) has six specimens. Four of these were registered sometime between 1900 and 1945, but collection dates are not available. Localities are Helensburg, Long Bay, Middle Harbour, and Coast Range at Botany Bay. The Middle Harbour specimen, at least, possibly originated from, or was known to, Krefft because this collection locality was specifically referred to by him (Krefft 1869). Two other specimens in NMV collected in 1975 are from Yal Wal (Nowra) and Royal NP. The Western Australian Museum (WAM) has three specimens, all from Woronora Dam in the 1960s and 70s. European collections In Germany, there are four museums with records of 13 H. bungaroides specimens. The Museum fiir Naturkunde, Berlin (ZMB) has eight specimens. Two of these were purchased from “Salmin”, a dealer in Hamburg who traded with Krefft. They are undated, but it is known from the Annual Reports that Krefft sent Salmin reptiles in 1866 (see Appendix 1). The ZMB also has three specimens labeled “Krefft” specifically. Another two specimens are from 1867 and donated by Richard Schomburgk (1811-1891). Schomburgk was Director of the Botanical Garden in Adelaide from 1865-1891. None of these seven specimens have specific point localities, i.e. either “Australia”, “New South Wales” or “Sydney”. The eighth specimen in the ZMB was donated by the Berlin Zoo on 12 September 1913, and the original collector and collection place are unknown. In the Zoologisches Museum, University of Hamburg (ZMH), there are two specimens: one from Krefft; the other with no collector details. These specimens are recorded as from “Sydney” and “Australia” respectively. The single specimen in the Senckenberg Natural History Museum, Frankfurt (SMF), from “eastern Australia” was donated in 1911 by “O. Frank”. We have no details on “O. Frank” or any other information on where he found his specimen. The Zoologische Staatssammlung, Munich (ZSM) had two H. bungaroides from “New South Wales” registered in 1920 and 1928, but these were destroyed during World War Two (D. Fuchs pers. comm.). Seven specimens were found in the UK. Six are preserved in the Natural History Museum, London (BMNH), and one in the Oxford University Museum of Natural History (OUM). One BMNH specimen was presented by the ‘Earl of Derby’ in 1847 (see also Ginther 1858; Boulenger 1896). This was Edward Smith Stanley (1775-1851), the 13th Earl of Derby. Two specimens in the BMNH derive from 1855. One of these was donated by the Zoological Society of London (ZSL), but the collector of this specimen is unknown. It was possibly John Gould, since he collected many specimens in Australia and also worked for ZSL. The second 1855 specimen is from the “collection of Captain Stokes”. This was John Lort Stokes, who was on the Beagle surveying expedition to Australia from 1837-1843. There is also a specimen in the BMNH registered 1859 that was presented by Proc. Linn. Soc. N.S.W., 130, 2009 J.M.HARRIS AND R.L. GOLDINGAY “Dr G. Bennett”. This was George Bennett, who was an early Curator of the Australian Museum from 1835-1841, and a Trustee of the Museum from 1853- 74. The other two BMNH specimens were purchased from Krefft and were registered on 16 June 1863. The only locality data with these specimens are “New South Wales” or “Australia”. The single specimen in the OUM was collected at “Sydney” by Francis Pascoe (1813 - 1893). Pascoe sailed to Australia in the Buffalo, captained by John Hindmarsh (first Governor of South Australia). After Pascoe’s death his large collection of zoological specimens was presented to the OUM by his daughter in 1909 (M. Nowak-Kemp pers. comm.). In France, the Muséum National d’Histoire Naturelle, Paris (MNHP) has four specimens. One of these (no. 7679) is the type of Alecto variegata (a junior synoymn for H. bungaroides), with locality given as “Australia”, collector/donor as Pierre Francois Kéraudren. This specimen is referred to by Schlegel (1837), Duméril et al. (1854), and Guibé and Roux-Estéve (1972). There is also a specimen from Port Jackson donated by “Quoy and Gaimard” (i.e. Jean René Constant Quoy and Paul Gaimard), collected some time prior to 1829 when these French naturalists visited Australia. The actual location data provided to us are for Middle Head. This specimen is also mentioned by Schlegel (1837), Duméril et al. (1854) and Guibé and Roux-Estéve (1972). Another MHNP specimen was collected from “Australia” by the French naturalist/specimen dealer Jules Pierre Verreaux some time in the early 1840s (also in Dumeril et al. 1854). According to the MNHP donations book, it was received in December 1846. The fourth MNHP specimen is a skull registered as no. 1991-4163. This specimen has no date, collector or locality details, but it is a different specimen to the above three, and it is believed to be from the same era, i.e. 19" Century (I. Ineich pers. comm.). In Austria, three specimens are preserved in the Museum of Natural History, Vienna (NMW). These are NMW 27699:1-3 and are dated between 1863 and 1877. There are no collector or donor names recorded with any of these, and the original label for these specimens indicates “West Australien” (=Western Australia). Photographs of the specimens supplied to the authors confirmed that the identifications are correct. However, the locality data is certainly erroneous. Other Australian snake specimens in the NMW collection were purchased from the dealer “Gerrard”, and it is possible that specimens with confused localities were sold by him, including these three H. bungaroides specimens. In Denmark, three H. bungaroides are preserved Proc. Linn. Soc. N.S.W., 130, 2009 in the collection of the Zoological Museum, University of Copenhagen (ZMUC). Two of these are dated 1862 and from “Sydney”, but no collector details are recorded for either specimen. The third from “Australia” was donated to ZMUC by “Dr Giinther” in 1867. In the Netherlands, the National Museum of Natural History, Leiden (RHNH), has five H. bungaroides specimens. One of these from “Nouv. Hollande” (Australia) was donated to RMNH by John Gould. Another two specimens recorded as from “Nouv. Hollande” are dated 1849 and were donated by “Frank”. This was probably G.A. Frank, a natural history dealer based in Amsterdam. A specimen from “Botany Head”, dated 1862, was received as a gift from the AM. The fifth specimen was also from the AM, but this has no date and no locality. Naturhistorisches Museum, Basel, Switzerland (NMB), has one specimen of H. bungaroides from “Australia”. It was donated in 1882 by Dr. Fritz Miller and is registered as no. 2188. Miiller apparently contributed many purchased or traded herpetological Specimens to the NMB in the years between 1880 and 1890 (R. Winkler pers. comm.). Advice received was that in this period, Miller worked voluntarily for the NMB and cared for the reptile, amphibian and fish collections. At the Zoological Museum, University of Liege, Belgium (MZULG), there are five H. bungaroides mentioned in the museum register. All arrived between 1856 and 1875 from specialised natural history shops (C. Michel pers. comm.). Three of these do not have localities, but two indicate “Melbourne”. If the latter two are truly H. bungaroides, then the recorded localities are also incorrect. However, as with the specimens from the NMW further study of MZULG specimens are also required to ascertain whether this is the case. In Italy, the Museo Civico di Storia Naturale, Genoa (MSNG), has one H. bungaroides (8687). The specimen was acquired in 1879 from the Godeffroy Museum of Hamburg (Germany), a private institution founded in 1860 by Johann Cesar Godeffroy (1831- 1885). The MSNG acquired specimens from the Godeffroy Museum by means of nine catalogues edited from 1864 till 1884 that listed duplicates put up for sale (G. Doria pers. comm.). No locality data are available for the specimen held at MSNG. North American collections In the US, there are five museums that together hold nine H. bungaroides specimens. The Field Museum of Natural History, Chicago (FMNH), has two specimens both collected at “Waterfall” in MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE the 1950s. One was collected by William Hosmer, a well-known herpetologist who worked as a field collector for the FMNH for many years and sold his Australian collection to that museum. It is known that the other FMNH specimen was collected by B Kaspiew, although we have no further information about this person. The Museum of Comparative Zoology, Harvard University (MCZ), has three specimens: one from “New South Wales”, received from Krefft in 1876; one from “Australia”, received from ““W. Keferstein” and registered in 1865; and one from “Gelle, Mt. Wilson, Blue Mountains”, received from the AM in 1914 (Loveridge 1934). The National Museum of Natural History, Smithsonian Institution (USNM), has a specimen catalogued in about 1872 with no locality details or collector name. The third from Sydney dated 1911 was received from “Julius Hurter”, a Swiss-American naturalist and early Curator of the St. Louis Academy of Sciences. The single specimen in the San Diego Natural History Museum (SDNHM) was originally sent there by the AM on exchange to Van Wallach and Richard Etheridge (San Diego State University) for Wallach’s studies on the visceral anatomy of the Australian Elapidae (see also Wallach 1985, 1998). A copy of the “specimen invoice form” shown to the authors was dated 19 January 1982 and indicates that this H. bungaroides was a “no data specimen”. The single specimen in the University of Illinois Museum of Natural History (UIMNH) has no location recorded with the specimen and was apparently “purchased from the AM” but the date for this transaction is unknown. It was originally catalogued into the very old zoology collection (<1943) and the Curator at the UIMNH suggested that it was probably from the 1920s judging by its very low “Z” catalogue number (006) (C. Philips pers. comm.) DISCUSSION Of the 159 specimens, accurate locality records were available for only 98 (62 %). The AM contributed 77 while another 25 institutions contributed the remaining 82. Several of the latter (detailed in the notes above) are highly significant: two records for Middle Head (dated <1829; 1935), two for Botany Bay (dated 1862; <1935), one for Long Bay (dated <1935), one for La Perouse (dated 1915), and three for Ku-ring-gai Chase NP (dated 1967). Four of the AM specimens (dated 1904/5) were from the same location at Long Bay as that above and two specimens (undated; 1895) were from the same locations at La Perouse as that above. Significant specimen records from the AM include those from the western side of the Blue Mountains (Bathurst: dated 1979; Ilford: dated ca. 1962), and from Mudgee (<1964). Other significant records are those from within the vicinity of Shoalhaven Formation geological outcropping along the western and north-western rim of the Sydney Geological Basin, the presumed limits of the species’ distribution. Whilst perhaps the species is absent there today, it gives a clear indication that some of this otherwise presumed habitat was in fact occupied by H. bungaroides. With many of these historical records collectors probably gave locations that covered wider districts or the specimens were allocated names of the centres they were brought to from the field. This is likely to be the case for the western records from Bathurst, north of Bathurst and Mudgee. The specimen locality data were mapped and contrasted with the 67 records in the Atlas of NSW Wildlife (Fig. 1). Two specimens from the AM (dated 1969) and one from SAMA (dated 1973) had as the locality data a site close to the location of the AM itself. We believe the co-ordinates for these three relatively recent specimens to be incorrect, and so excluded them from the map. The distribution of the museum records shows some concordance with the Atlas data. Both databases show aggregations of records in the Katoomba (Blue Mountains), Waterfall-Heathcote and Nowra (Shoalhaven) areas. Surprisingly, 37% of the records in the museum database are from Royal NP (28) and the adjoining Heathcote NP (8) and Garrawarra SRA (1). One location in Royal NP covering an area with a radius of 2 km contributed 23 specimens with collection dates spanning 1951- 72. These observations identify and confirm the currently known ‘hotspots’ of the distribution. We can also contrast Figure 1 with the only map previously published based on Australian Museum holdings (Longmore 1986; 50 specimens). There are about 15 museum records since 1986 including several for the Blue Mountains area (including Wollemi NP). Including these on our finer detail map gives it greater completeness as it includes Atlas records and non- AM museum records. Hoplocephalus bungaroides is reported from only a small geographic area, as evidenced from the locality data available from museum specimens (Fig. 1). Krefft (1869) reported H. bungaroides from Port Jackson, Botany Bay, Middle Harbour, Lane Cove and Parramatta, although as pointed out by Cogger et al. (1993), there have not been records from these areas for quite some time. These data indicate that the only museum specimen from Port Jackson was collected prior to 1829 by Quoy and Gaimard (MNHP 7678). Proc. Linn. Soc. N.S.W., 130, 2009 J.M.HARRIS AND R.L. GOLDINGAY Cessnock e llford » A ++ + + A Bathurst se Colo Heights *A~ th quithgow Records A museum a ee Kilometres + Wildlife atlas Fa 0 10 20 30 40 Map produced by Greg Luker, SCU GIS Lab, 22/8/2006 Figure 1. Geographic distribution of Hoplocephalus bungaroides as indicated by museum records and records in the Atlas of NSW Wildlife. Proc. Linn. Soc. N.S.W., 130, 2009 7 MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE At Botany Head, a specimen was collected in 1862, and ended up in Leiden, Netherlands, sent there by the AM (i.e. Krefft). There is also an AM specimen from Botany dated 1909 and another in NMV undated, but registered some time between 1900 and 1935. Middle Harbour museum specimens are in the NMV and MNHBP. It is likely that the Botany Bay and Middle Harbour specimens were known to Krefft, because these localities were specifically referred to by him (Krefft 1869). Of the 159 H. bungaroides specimens located, none had locality details recorded as Lane Cove or Parramatta. Thus, Krefft knew of H. bungaroides records from these locations, but it is uncertain whether he collected specimens from there. Krefft did undertake snake collecting in many places in the vicinity of Sydney. Rose Bay, Randwick, Manly, Coogee and Middle Harbour were reportedly principal localities (see correspondence between Krefft and Giinther in the archives of the AM). The annual reports of the AM are unequivocal in reporting that H. bungaroides specimens were sent to the Civic Museum, Milan (Italy), in 1865; R.H. Beddome (India) in 1867; and Berlin Museum (Germany) in 1871 (Appendix 1). The Exchange Register also indicates that one H. bungaroides was sent to the Madras Museum (now Government Museum, Chennai). In relation to the first of these, we made enquires with the museum in Milan (MSNM; Appendix 2), but H. bungaroides could not be found on the shelves or in the collections register. However, we found an H. bungaroides in Genoa, Italy (MSNG; Table 1), but this is dated 1879, and it is unknown whether this snake arrived at MSNG via the AM. In relation to Beddome, it is known that he was a naturalist and a British military officer posted to India. His zoological collection together with that of his son-in-law (G. C. Leman) was sold in 1935, and much of this material is now in the National Museums of Scotland (NMS); National Museum of Wales (NMW); and the Natural History Museum, London (BMNH). However, only the latter institution has H. bungaroides represented, and these specimens are all dated prior to 1863. Hence, the fate of the AM’s 1867 specimen sent to India is also unknown. The AM 4. bungaroides sent to Berlin in 1871 are still preserved in the ZMB. This museum has three H. bungaroides from the AM (Krefft), and another five specimens that arrived via other avenues. Unfortunately we were unable to confirm the presence or absence of H. bungaroides at the Government Museum, Chennai, because no advice was received in reply to our correspondence. The annual reports of the AM were quite vague in terms of the reptiles sent to de Robbillard in Mauritius; dealers Puls, Salmin, Fri¢ and Damon; and museums in Hamburg, Leiden, Madras, Paris, Madrid, London and Harvard (Appendix 1). Of these, we managed to track down H. bungaroides specimens collected/ donated by Krefft in Hamburg (ZMH), Leiden (RMNH), London (BMNH) and Harvard (MCZ). We can also confirm that Salmin received some H. bungaroides specimens (presumably from Krefft) because two from him were located in Berlin (ZMB). We found no evidence that other high-profile dealers such as Fri¢€ (Reiling and Spunarova 2005) received H. bungaroides from Krefft or anyone else. This review demonstrates the value of museum specimens as a source of information on species’ distribution (see also Shaffer et al. 1998). It’s widely known that much Australian material has made its way to 19" Century collections overseas, but the details of such holdings are still not easily accessible and so our contribution at least makes such distributional information available for H. bungaroides. Collectively, the museum data show specific records for Sydney’s urban areas - Botany Head, La Perouse, Long Bay, Botany, Concord West, Randwick, Middle Harbour and Port Jackson. These localities represent part of this species’ historical geographic range that has now been eliminated (see also Swan et al. 2004; Shine et al. 1998). Increasing our understanding of the historic distribution of H. bungaroides is of considerable importance because continued habitat clearing and fragmentation may eliminate this species from an area and without an historic record may lead to disagreement about whether an area is actually suitable for this species. For example, Hoser (1995) categorically refutes that H. bungaroides occurred in Ku-ring-gai Chase NP but three H. bungaroides specimens in the SAMA have collection details dated 1967 for that locality and there is no reason to doubt their authenticity. Recent surveys there (1998/9) failed to detect H. bungaroides (Newell and Goldingay 2005), suggesting it may now be locally extinct. The museum specimen localities provide a focus for increasing our understanding of the geographic range of A. bungaroides. There are three broad areas with aggregations of records: Katoomba (Blue Mountains), Waterfall-Heathcote and Nowra (Shoalhaven area). These areas also show aggregations of records in the Atlas of New South Wales Wildlife (Fig. 1). These may represent areas of highly suitable habitat for H. bungaroides. However, there is likely to be collecting bias evident with these data. For example, a few areas near Waterfall contribute 37% of all specimen locations, though records span a 27- year period. Recent detailed surveys in Royal NP (i.e. Proc. Linn. Soc. N.S.W., 130, 2009 J.M.HARRIS AND R.L. GOLDINGAY Waterfall) indicate that H. bungaroides is uncommon there (Goldingay 1998; Goldingay and Newell 2000; Newell and Goldingay 2005; Goldingay and Newell unpubl. data). The failure to detect H. bungaroides in recent surveys of national parks surrounding the Hawkesbury River where there are few historic records (Newell and Goldingay 2005) suggests that the species’ distribution is much more patchy than what might be predicted based on the presence of apparently suitable sandstone habitats. Further surveys of suitable habitat in areas without records need to be conducted. Records in the north-west of the species’ range (Bathurst: dated 1979; Ilford: ca. 1962) also highlight areas where further surveys need to be conducted. These represent the most western records of the species and a population in this area may show some genetic divergence and be of considerable conservation significance. The identification of museum holdings of H. bungaroides may be useful for a range of future research studies. This includes morphological research and further descriptions of diet based on stomach content analysis (e.g. Shine 198la,b, 1983; Keogh 1999). Furthermore, these specimens may provide a source of tissue samples for genetic studies that could contribute to an understanding of whether H. bungaroides has lost genetic diversity over time or if unique genotypes have been lost (see also Keogh 1998; Slowinski and Keogh 2000). Our collation here provides a record that will facilitate the use of specimens in this way. ACKNOWLEDGEMENTS We are grateful to the many curators and collection managers that responded to our enquiries. For data and information on museum holdings of H. bungaroides we acknowledge Ross Sadlier (AM), Russell Graham (ANU), John Wombey (ANWC), Colin McCarthy (BMNH), Maureen Kearney (FMNH), Paul Daughty (SAM), Carolyn Secombe (SAMA), Jose Rosado (MCZ), Raffael Winkler (MHNB), Stuart Norrington (MMUS), Ivan Ineich (MNHP), Giuliano Doria (MSNG), Nicole Kearney (MV), Christian Michel (MZULG), Marie-Dominique Wandhammer (MZUS), Franz Tiedemann (NMW), Paul Horner (NTM), Malgosia Nowak-Kemp (OUM), Andrew Amey (QM), Koos van Egmond (RHNH), Bradford Hollingsworth (SDNHM), Jens Kopelke (SMF), Chris Phillips (UIMNH), Ken Tighe (USNM), Rainer Guenther (ZMB), Alexander Haas (ZMH), Mogens Andersen (ZMUC) and Dieter Fuchs (ZSM). We thank Leoné Lemmer (Australian Museum Library) and Glenn Shea (University of Sydney) for assistance with identifying relevant literature, and for advice on Australian material held in overseas institutions. Further help was received from Van Wallach and Richard Etheridge. Stephen Sleightholme provided contact details Proc. Linn. Soc. N.S.W., 130, 2009 for some European museum curators, and Margaret Pembroke (Southern Cross University Library) tracked down some old literature. The map of distribution records was produced by Greg Luker (Southern Cross University GIS Lab). We thank David Newell and Ross Wellington as well as several anonymous referees for comments on an earlier draft of the manuscript. REFERENCES Boulenger, G.A. (1896). Hoplocephalus. In “Catalogue of the snakes in the British Museum (Natural History). Volume III., containing the Colubridae (Opisthoglyphae and Proteroglyphae), Amblycephalidae, and Viperidae’. Pp: 348-350. Cogger, H.G. (2000). Reptiles and Amphibians of Australia. 6" edition. Reed New Holland: Sydney. Cogger, H.G., Cameron, E.E., Sadlier, R.A., and Eggler, P. (1993). The action plan for Australian reptiles. Australian Nature Conservation Agency, Canberra, Australian Capital Territory. 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RIL] PT Op oasnj . UOAT “O[[OANJLNY OITO}SIFY,P UUNasNyY - So[PINJLN SPIOUSTD op OUTZUSSIV Oasnyy - UNINY “o][oINJVNY IITOSTFY.p WNasnyy a O[[IT [ens] uoloepuny ee d[QOUSIDH ‘o][o1NjeNY 9.110}SIF] Pp Unasnyy vujuos1y - uoliq ‘ssousI9g seq s}[novy ‘onbisojo0ozZ o110je10ge 7] - Kauphs Jo AjisioAtup) “winesnyy ASojoo7Z |[oMsey] - AoueBN ‘Saousldg sad 9}[Novy BI Op d130]007 ap omO}eI10gGeT é PIURUUSeY] JO A}ISIOAIUL) ‘UOT|DaT[OD AdoO[o0Z douRAy 2 ABOTOUYSST, JO 9jINjSUT ouINogey] [eAOY ‘UOIaT[OD ABojooZ “ NYAIN] JO AISIOAU) “wuNesny] [eoIs0;ooZ ‘: SUINOGIaJ\ JO AIsIOATUL) “WINaSNy] SSOLy, ¢ uinasny] AVIsIoATUp) B[AYSeAAL * puvjsucon() Jo ApisioAtup) ‘uinesnyy As0;007 2 ArOjSIF{ [RANeNY JO UNasnyy YsTUULF p puejsuq MoN JO AjIsioAtUy) ‘uunasnyp Ad0;007 puryluny P peqoy ‘Alo[eH jy pue winosny] ueruewse], = ArO\STF [RANJeNY JO Unosny] ULIUO}S_ n uojsoouney ‘Alo[eH iy pue winesnyl VIIO}DIA UdONd = Ipuelfi, ‘puelfi, Jo winosny\ ES AjIsIOATUP) o1renboryy “winasny] Soouatog [BoIsOO1g vIUuO}Sy ul[e.ysny u uonnjysuy u uonnjyysUy ysonboi uo UvYy} 9.101 331dsop Ajdo. jou pip (-) & Y{IAA SUOTNINSUT SUIMISNU ALoY) UT Poy 910A SUdUIDOdS saplossung “FZ OU VY) pordos (,.) & VIA poyxAvUl SUOINIHSUT :930N7 SA[day =y ‘auIeU UONIHSUI Vey) pue A1QUNOD Aq ATTVOHequvydye uo) IsAy LIPEUSNW IIA post] o1v Solu” ‘soUdpuOdsaa.109 [fvula Ano 0} Ajdo. jou pip 10 saplosnsung snjvydasojdopyy ay[eus popeoy-pvo.1g 94} JO SUdWAIDIdS P[oY jOU Op JE) SvaSADAO puUL LITV.QSNY Ul SUOLNIWSU] :7 xIpuoddy N — J.M.HARRIS AND R.L. GOLDINGAY ee ne eh sooualos Jo AWopeoy uvIssny ‘UIMasny] [ed150[007 AUISIOATUL) 9321S MODSO/| “LUNESNA] [29130[007 MOODSOJ] “WINISHA] UIMIE 3121S RISSHY “WUINEsNy] [LOISO[OOZ ueLIEqIS A}ISIOATUP) AOWIeUY ‘ATOYSTY] [eINjeNY JO wnesny| vISsny AjISIOAIU) 1eATOG-soqeg “UINasny] [ed1s0[00Z jsoleyong “winesnyy A10}S1H [RANIeNY NIQES “wunesnyy [eyUeyNIg BIULULOY RIQUILOD “UIMasny] AIO}SIF] [eINIEN RIQUILOD JO AjISIOATUL) ‘OISOTOdoONUY 2 Od1SO[00Z nasnyy UOgsTT JO AjISIOATUA) ‘[eINIeNY VIIO}SIEY Op [eUOIORN] Nasnyy pllopeyy ‘[eyouny op jediorunyy nosnyy OVO JO AYISIOAIUA) ‘[RINJeN VIIOISIEY op nosnyy UOgsTT ‘eIZO[00Z ap oUED [esnj}.10g Moyer “ApISIOAUP) URIUO][aISeL “LUNOSN] [RdISO;OOZ saouaIag Jo AWopeoy Ysijodg MP[IOIN JO AjISIOATUL) “UMNasNyy AIO\STH] [RANIeNY MesIeA ‘ASO[OOZ JO aN NSU] pure winesny[ puvlod O]SE JO AjisioATU) ‘UNMasnyy ASo;OoZ udsIog JO AjISIOATUP) “UOT}Oa]]0D ASo[007 JOBULARIS “SUI[OPAV YSISO[OOZ wnasnyj] JosUeAL}S wleypuory ‘Asojooeyory pue Aro}sip{ [eINyeN JO Winosnyy AGMAON Ulpounq “winasnyl 03210 pueppny ‘wnesny] pueppony yomnyoysiyO “winasnyy Aingiayued puelesZ Mon Wepio}suly “UINesny] [2913 0]007 ION JO AjISIOATUP) ‘UOT}IET[OD [eoIso[007Z WepioyOY ‘WepsoyOY WinosnuInNjeN, JYOIMseey] Wines] YOSOJsTYINNyeNy SpuLloyjoN SINOGUIOXN] ‘a][oINJeN SI1O}ST_Y pp [eUONeN sosnyy s.moquiexn] RSIY “VIA\eT JO wnesny] ATO}sI_] [eINjEN RIAL] OAYO], “Winasny] USING [eUOeN| “UOT}Oa][0D AS0[007 AIO}SIF [eINjeN JO Uinosnyy] eyesCO AUISIOAIUL) NYOYOL, “A1O}STF] [INN FO wNasnyy umnesny] ArO}sIP jernjeny NAsNAYLIDY uvder He ERP RE AUSIOAU) Wel[se_ “UONd9T[0D [eo1so[00Z pIssOy ‘ALOJSIF{ [PINJeN| JO WNosny [LLOUIAOI OUI, “AIOjSIH{ [RANWRNY FO winesnyy so[den Jo AjIsIaAUp) ‘Oo1s 0,007 Oosny| OULIA[e IP RIISIOAIUL) “ODISO[OOZ Oesny/| BUSI “OD1SO[O0Z Oasny/| ULIN], ‘einen SZUIIOG Ip sjeuoIsay Oosny/ I[VINJON YZUSIOG Ip s[BUOISOY Oosny/y eZUatdes eT, sWIOY JO AyISIOATUL) “VISO[OOZ Ip Oosny/y RUSO[Og ‘CUBO]O IP RUISIOAIUL) ‘eISO[OOZ Ip Oasny/[ vAOpE_ JO AjISIOAIUL] “eISO[OOZ Ip ossny/| eUSO]Og ‘voneAlas vuney vy Jod oyeuoizeN] O1NI1SU] [op Oosny| BUOIOA “OTRINJLN] VIIOJS IP OSIATD oasny| AUDA “A[LINJLN VIO IP OSIATD Oosny[ ULIIP] ‘O[LINIVN VIIO}S Ip OOIATD Oesny| OISOLL], Ip BLINN] VIIOJS IP OIA Oasnyjp| BSIq JO APISIOAIUL) ‘OLIOPIMIAT, [op 9 oTRANRN VIIO}S Ip Oosnyy SOUdIO][Yy ‘WInesny] AIO}STF] [LINJeN SOUSIO].Y sWIOY ‘ABO[OOZ JO winesnyy STATO RIOUTLD “AIOISIF] [RANVeN] OJUATVS oy} JO WNasnyy STATO ArO}SIF{ [PINJVN JO WNasny] IIAID OIe}OIO}UOP] OAONUTESeD leg Jo AjIsIoAluy) “WInesny] [eoIsoO]0OZ leg ulfgnd eset[oD Aur Uljqng “winesnyy ArO}sT [eINeN Alea] puvyjory TeqyeA ‘AISIOAUp) eIypuy “Wnasnyl [291d0[007 ByNO[VO “winesny] UeIpuy TeuUUSYD “Winesny] JUSWUUIOAODH) UOI9I[OD eIpuy Jo AoAINS [eoIso;007 IY] MON ‘ArO}STH [eINIeN JO wuNnesnypy [eUoneN Aequiog “uoysa][0D Ajeto0g Arojsipy [eAnjeN Avquiog jsodepng ‘uinesnyy ArojsIP jenjen Ueesuny, vipuy Axesunyy suo JO AyIsIoATU) “WInasny/] [eOIsO[00Z 999015) UdSUIQNY, JVjISIOATUP] “wuNesnyA] ASO[OOZ UWINasny] SOYOSIUIZIPSULIEL], pun Soyssiso[007 AUISIOALUP) JOUINT-UNIW] “WNyWSUy SoyosIs0[007, PIPMSHIOID “UmMasnyj] UsyOsIso[007 SINQoploH JO AjIsIOATU) “WUINssny] [POIsO[OOZ ussUIOH “UINasnyj [eO1s0;007 ARISIOATU)-S}YOoIQ] V-UeYSIIYO “Wnesny] [2d1d0[00Z7 YOO}JSOY JO AjISIOATUL) “WOT}D9][OD [eoIs0;007, ponunuos Aueu1es5 13 Proc. Linn. Soc. N.S.W., 130, 2009 MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE Ca SS RG RS DS ete CD SC Cs * i De ue OE AUISIOAIUA) SUNOA WeYysiig “WInasny/A] SOUSIDS oFIT QOUSIMET “UINasnyAy AIOISIFY [BINJeN] AjISIOATUE), Sesuey eUvIpUy ‘osoT[OD weypieg “winasnyy a100;] Ydasor AIO\SIH [RANJeN] JO wunasnyy OYepy JUIN [eINeN JO Winasny] UO\sNo}{ BIUAOFIeS ‘A1Ojsip [eanjyeN JO wnasnyy AeTpeA e215) Aro\sTpY [eINeN JO winasny] eIs1090 Alo\sipy{ [eANJeN FO wNasny] epr10] 4 ByuepYy ‘AIO}SIF{ [eINjeN JO Winesnyy yUequis, AIO}SIF{ [RANLN FO wNesny seT[eq SoJVIGO}IOA JO Winasny] ATISIOATUP) [[aUI0D AlO\stP [RINBN JO winasnyy puejars[D BUI[OIRD YNOS “UINesny] UO\sapIeYyD AVISIOAIUL) 9381S UOJBUTYSEA “UUINEsN] IouUOD ~yY septeyo AlO\SIF [VANJRN JO WINasny] sIdauIeD uoNdaT[0D Asojojodiayy sous Jo Awiopeoy erusojeD SIOUN]]] ‘PIOF yoo ‘A1OjsrY [eANyeN Jo wunasnyy coding aInj[ND pue ArojsTFY [enjeN] JO wWinesnyy syINg neMey “winesnyy doystg vjosouUIPy FO AjIsIOATUP) ‘ArO}STHY [eINWeNY FO wnasny] [[9q winesnyy ATISIOAIUp) 93%19 Avag uUNSny UOdIT]OD Asopojodiay] AjISIOAIU 9}e1g PUOZIIY YOK MON ‘AIO}STH [RANWeN JO winosnyy URoTIowy AIOYsTH [RANJRNY JO wnasnyy Bureqeil Vy vIydjape]iy_ ‘Soousiog jeinjeN Jo Awoperoy BILIOULY JO $9}¥}S poy AjISIOAU) Usapleqgy ‘uInesny] ASO[007Z Spoo7T JO AjISIOAIUA) “UOTJD9][O_D [eo1s0[007 Alay[eD iy pue umnesnyy AqID 19}S9910/\\ wunasny] AS0[007 sepunqd Jo AjIsIOATU) espliquieD ‘Aso[0oZ Jo umnasnyy AjIsIoATUy) pURlol] WIOYION UI wuNasnyy 10}S|) xossq “Uinosny\] Uoprep ONES Jojoxq ‘Winasny] [RWOWel Weg py [eAoy SdIAIOS Winasnyy SUIPRoYy SOIAIOG Splosoy pure sunasnyy AID YNowsI0g yynouwrAyg ‘AlayjeH iV pue wunesnyy AyD ynowA,g YsInquipy JO AjIsIOAUP) ‘UOMIET[OD A10\sTH [eINIeNY Ysinquipg ‘pueyjoog Jo sumnesny] [euOHeN Jjipred ‘soyeAA JO unasnyy [euoeN, Jojsoyoury] JO AjISIOAIUL) “WINasnyy Jo]soyouRy/\ suinesny] [OOdISATT uopuo7T ‘Aja100g ueouurT spoe7 ‘winasnyy AID spose UOpuoT “asaT]OD ssury [epucy ‘winoesny] [epuoyy SITYSI9]S9OMO]H “UINesnyA] opisAUNOD s1Oo/] UYOL BS AP DS PS * mre a ED aD BY yA Be SP SD YoIMsdy] “umnasnyy yorMsdy uOpuUOT ‘suossding Jo dda][0_D [eAOY ou} Je UNosNyy URLIO}UNFy PpuRpoos “‘Mosse[H ‘Alol]eH jLy pue winesnyy uewojunyy UOpUOT “wnssnyl UeWTUIO HY Alol[eD ily pue winesny] projoiay a[ISLOMAN “WUNasny] Yooouryy JaJSOYOUI\ “AdIAIAS sunesnyy AjuNOD ostysdurey UOPUOT B8a]]OD AjisioAtuE) “ASO;OOZ Jo winasnyy jUeIH aysy1eg “ASo;ooZ Jo wuinasnyy aJoD [oIslIg ‘AlaljeD yy pue umesnyy AyD [oIsg uoJYSIIg “AIOSI [eINyeN Fo unosnyy YOog (Ja]soyouRy]) UOWOg “sumMesny] UoO}TOg SMOIPUY 1§ JO AISIOATUL) “LUMasny] MoISINOg [[9q wopsury paytuy WINYJOTOS “WnosnuiNjeN Spuo,-ep-xneyD eT ‘a]joinjeN S110]sI}{ Pp sesny/] quuRsney ‘s180[007 ap sasny| Ug[eH 3S “uinesnuinyen] YoInZ JO AjISISAIUL) “WINesny [29150007 SINOQILY “WINesnyy AIO}STH [eANyeN Wlog “winasnyy A1O}SIP] jenjen, PADUDD “o][OINJEN SI1O}STE].p wunosny/y Ld REYAL AWS B.10Q9}0H “UInasnyy A1OjSIE] [eANJeN] 3810qQ9}0H AjISIOAIU) puny “wuiMesny] [RoIs0;00Z A1OjSIH{ [ene JO winasny] YstpaMms Wapaas PUISIO}VMIIA\ JO AjISIOATUE) “UMESNy] ASO[00Z Yyosoquay[ars JO AjIsIoATUP) “WuMesny] [2d1d0[00Z7 RUO}JOIg “WINasny] [BeASURIT, UMO], odeD “wnesny] ULOLIZYy yNOS winesny] Woqezi[q 0d UloJUOJWISO] g “WINesny] [PUOHeN SINQZILULIO}O1 J “WNesny [PIVEN Ag]loquiry “winasnyy 1OSs1g 9; ueqing “umesnyy ueging UMO}SWeYeIDH “wuMosnyy] Aueqyy BILIFY YINOS euelignly ‘A1ojsip [eINjeN] JO Winesny] URIUSAOTS BIUAAOTS Aofepieg “winesny] dyssuesg EPIEAOTS PLIpey ‘So[einjeN SeloualD op [euOIoRN Oasnyy BuoIOIeg “BOISO[OOZ Ip Oasnj| winesny] A10}STF] [VINJEN ,.SPISO[S] SINT, osuving ‘evoosny] UdIZ}USIZ INjeN OxEpyesuesng ulvds penuguos z xipueddy Proc. Linn. Soc. N.S.W., 130, 2009 14 J.M.HARRIS AND R.L. GOLDINGAY SEO OE BX ES I Ee EE * * AIOMSIFT [LINJLN JO UMosNyA] UOSOIO Jo AYISIOATUL) WINasny] 2101S BYSeAQON JO AjISIOATUP) uljsny ‘sexe] JO AjISIOATUL) ‘SUONOOT[OD ArO}SIH [eANJeN] sexo, AIOJSIFT [L.INILN JO Urnosnyy BIeqIeg vIULS ALOJSIFT [LINILN JO UMosny] VUOYLPO [GON Weg AjIsIoATU) a[eA ‘AIOISTEY [RAMJLNY JO tunesnyy Apoqeog umMasnyl AjISIOATU) 21219 USA SOOUSIOS [LINN JO Winasny] 97k}S BUTTOARD YON uinesnyl 972} JOA MON AIO\SIH [BANILN JO UNosny] OOTXA|A] MON styduroyy JO AyisIoAtuy) ‘Asojooz Jo umosnyy] OOIXO] MON JO “AlUP) ‘ASO[OIg Wo}soMUINOG Fo winesnyj| wnasnyy 91]qnd SaxNeAyAl ponuUods voLeUry JO Sojev}S pou, 15 Proc. Linn. Soc. N.S.W., 130, 2009 MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE Appendix 3: Hoplocephalus bungaroides specimens reportedly held in Australian and overseas museums. Note: the authors have not personally confirmed the identification of any of these specimens by examination. Records are arranged alphabetically by museum abbreviation (see Table 1) then numerically by registration number. Abbrevia- tions: Coll. = Collected; Confis. = Confiscated; Don. = Donated; NP = National Park; NSW = New South Wales; QNPWS = Queensland National Parks and Wildlife Service; ZSL = Zoological Society of London. Collection Date Museum Rego. No. Locality details Other details = AM R 1440 - Registered 30/08/1893 - AM R 1603 La Perouse - = AM R 1722 La Perouse Registered 14/04/1895 6/04/1900 AM R 2696 Mount Wilson Registered 10/05/1977 11/10/1904 AM R 3646 Long Bay Registered 15/05/1977 12 Apr 1905 AM R 3675 Long Bay Registered 15/05/1977 28/04/1905 AM R 3678 Long Bay Registered 15/05/1977 26/11/1905 AM R 3847 Long Bay Registered 18/05/1977 26/11/1905 AM R 3848 Long bay Registered 18/05/1977 16 Dec 1909 AM R 4619 Botany Registered 22/05/1977 - AM R 11179 Randwick Registered /04/1934 1/11/1959 AM R 15676 Waterfall Registered 27/11/1959 Aug 1959 AM R 18939 Waterfall Registered 30/11/1962 - AM R 18940 Waterfall Registered 30/11/1962 Apr 1962 AM R 18941 Mount Keira Registered 30/11/1962 - AM R 18942 Waterfall Registered 30/11/1962 - AM R 18943 Waterfall Registered 30/11/1962 - AM R 18944 Waterfall Registered 30/11/1962 - AM R 18945 Waterfall Registered 30/11/1962 - AM R 18946 Waterfall Registered 30/11/1962 - AM R 18947 Waterfall Registered 30/11/1962 = AM R 21071 Mudgee Registered 6/02/1964 2 Mar 1964 AM R 21219 Concord West Registered 6/03/1964 Feb 1969 AM R 30345 Springwood Registered 1/03/1971 8/09/1973 AM R 40309 Darkes Forest Registered 9/10/1973 2 May 1970 AM R 47415 Waterfall Registered 25/06/1975 22/10/1967 AM R 70034 Woodford Registered 1/02/1978 1966 AM R 74276 Royal NP Registered 16/06/1978 1966 AM R 74277 Royal NP Registered 16/06/1978 1971 AM R 74278 Waterfall Registered 16/06/1978 1971 AM R 74279 Sydney Registered 16/06/1978 1969 AM R 74280 Nowra Registered 16/06/1978 1969 AM R 74281 Sydney Registered 16/06/1978 1970 AM R 74282 Appin Registered 16/06/1978 1972 AM R 74283 Waterfall Registered 16/06/1978 Apr 1972 AM R 74284 Woronora Dam Registered 16/06/1978 2 Oct 1972 AM R 74285 Nowra Registered 16/06/1978 2 Oct 1972 AM R 74286 Nowra Registered 16/06/1978 2 Oct 1972 AM R 74287 Nowra Registered 16/06/1978 16 Proc. Linn. Soc. N.S.W., 130, 2009 2 Oct 1972 2 Oct 1972 2 Oct 1972 1967 1968 1969 197] 1971 1969 Oct 1978 1969 5 Aug 1978 2 Sep 1951 Jun 1963 5 Sep 1980 1979 17 Oct 1986 9 Feb 1996 1/01/1996 Jan 1980 Aug 1992 Aug 1992 Feb 1998 ~1963-1964 ~1978-1980 ~1978-1980 ~1855 ~1847 Proc. Linn. Soc. N.S.W., 130, 2009 > = > < An a A A ah Ss SS S Ss S > < zZ > PP Ss Ss zZ J.M.HARRIS AND R.L. GOLDINGAY R 74288 R 74289 R 74290 R 74291 R 74292 R 74293 R 74294 R 74295 R 74296 R 76338 R 82584 R 84381 R 92955 R 103159 R 103162 R 103711 R 107684 R 107685 R 107716 R 107717 R 107718 R 107719 R 107720 R 118644 R 125335 R 125414 R 128548 R 131075 R 131143 R 131144 R 131145 R 144614 R 144720 R 144876 R 147417 R 147418 R 150348 R 151978 RO1868 R0O5040 RO5041 1855.8.25.?? 1847.7.29.40 Nowra Nowra Nowra Helensburgh Royal NP Royal NP Waterfall Waterfall Sydney Area Colo Waterfall Colo Heights Waterfall Heathcote Mount Macleod Morgan Woronora Dam Bundeena Bundeena Stanwell Park Woronora Dam Waterfall or Heathcote Waterfall or Heathcote ~ 15km NE Bathurst on Road to Sofala Sydney Evans Lookout, Blue Mountains Hazelbrook, Terrace Falls Reserve Kangaroo Valley Captivity Linden, Glossop Road, Blue Mountains Linden, Glossop Rd., Blue Mountains Sydney Wollemi NP Waterfall Tiajuara Falls Australia Australia; Presented: Earl of Derby Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 16/06/1978 Registered 30/10/1978 Registered 31/05/1979 Registered 14/05/1980 Registered 28/10/1981 Registered 28/10/1981 Registered 25/12/1981 Registered 7/04/1983 Registered 7/04/1983 Registered 7/04/1983 Registered 7/04/1983 Registered 7/04/1983 Registered 7/04/1983 Registered 7/04/1983 Registered 30/05/1986 Registered 28/03/1988 Registered 18/04/1988 Registered 31/12/1987 Registered 17/05/1988 Registered 19/05/1988 Registered 19/05/1988 Registered 19/05/1988 Registered 10/05/1996 Registered 15/05/1996 Registered 10/05/1995 Registered 10/05/1995 Registered 24/02/1998 Coll. H. Cogger Coll. Greg Mengden Capt. Stokes Collection Macgillivray collection ld ~1863 ~1863 ~1855 ~1859 ~1953-1956 6 Oct 1951 1876 1865 1914 <1837 Dec 1846 Jun 1836 1856-1875 1856-1875 1856-1875 1856-1875 1856-1875 1893 1893 1882 2 Aug 1863 12 Feb 1869 1877 ~1970s 9 Mar 1975 25 Nov 1972 1 Aug 1978 <1909 6 Jan 1959 30 Apr 88 4 Sep 1989 Aug 1862 No date 18 MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE BMNH 1863.6.16.50 BMNH 1863.6.16.55 BMNH 1855.10.16.109 BMNH 1859.6.30.10 FMNH 75118 FMNH 97310 MCZ R2525 MCZ R3642 MCZ R10282 MMUS RO501a MMUS ROSO1b1 MMUS RO501b2 MNHP 1991-4163 MNHP 3301 MNHP 7678 MNHP 7679 MSNG 8687 NMV D 4270 NMV D 4704 NMV D 51865 NMV D 51866 NMV D 65041 NMV R 12709 MZULG D.R.1883 MZULG R.E. 2657a MZULG R.E. 2657b MZULG R.E. 2657¢ MZULG R.E. 4221 MZUS 626 MZUS 627 NMB 2188 NMW 27699:1 NMW 27699:2 NMW 27699:3 NTM R1212 NTM R958 NTM R1115 NTM R1217 OUM OUM 4641 QM J52877 QM 347924 QM J49761 QM J61008 RMNH RMNH 1141 RMNH RMNH 1142 NSW; Purchased from: G. Krefft Australia Australia Australia; Presented: Dr G. Bennett Waterfall Waterfall New South Wales; received Nov 1870 Australia; Gelle, Mt. Wilson, Blue Mountains Mount Wilson coast near Sydney coast near Sydney Australia Port Jackson Australia - - type of Alecto variegata Australia Long Bay Middle Harbour Yal Wal, Nowra Royal NP Helensburg Coast Range, Botany Bay Melbourne Melbourne Queensland Queensland Australia Original Label “ West Australien” Original Label “ West Australien” Original Label “ West Australien” Jarra Fall, Nowra Woronora Dam Heathcote Woronora Dam Sydney Waterfall Nowra Botany Head, Sydney Museum Sydney Proc. Linn. Soc. N.S.W., 130, 2009 Registered 16 Jun 1863 Krefft ZSL B Kaspiew W. Hosmer Krefft W. Keferstein AM J. Verreaux Quoy and Gaimard Keraudren Acquired 1879 Registered 1900-1935 Registered 1900-1935 Registered 1900-1935 Registered 1900-1935 Rolle Rolle Fritz Miiller Coll. Graeme Gow Coll. Graeme Gow Coll. Graeme Gow Coll. Graeme Gow Coll. F.P.Pascoe Found under rock Captive specimen Confis. by QNPWS Captive bred gift of AM J.M.HARRIS AND R.L. GOLDINGAY - RMNH RMNH 1335 “Nouv. Hollande” (Australia) Gould 1849 RMNH RMNH 1336a “Nouv. Hollande” (Australia) Frank 1849 RMNH RMNH 1336b “Nouv. Hollande” (Australia) Frank 2/06/1915 SAMA R00463 La Perouse Don. AM; now missing 1967 SAMA R12099 Kuringai Chase W. Irvine 1967 SAMA R12100 Kuringai Chase W. Irvine 1967 SAMA R12101 Kuringai Chase W. Irvine 2/07/1971 SAMA R13433 Woronora River H. Ehmann Sep-73 SAMA R14116 Sydney G.N. Coombe 1980s SDNHM 63864 - sent on exchange by AM 1911 SMF 20532 eastern Australia Don. O. Frank ~1920s UIMNH 95151 - purchased from AM <1872 USNM 8050 - Catalogued about 1872 1911 USNM 56166 Sydney Coll. Julius Hurter 8 Aug 1964 WAM R53761 Woronora Dam G.F. Gow 9 Mar 1975 WAM R53762 Woronora Dam G.F. Gow 9 Mar 1975 WAM R53763 Woronora Dam G.F. Gow 1860s-1870s ZMB 4443 Sydney dealer Salmin 1860s-1870s ZMB 4444 Sydney dealer Salmin 1860s-1870s ZMB 5208 NSW Krefft 13 Sep 1913 ZMB 63510 Donated by Berlin Zoo - 1860s-1870s ZMB 63755 Australia Krefft 1860s-1870s ZMB 63756 Australia Krefft 1867 ZMB 63757 Recorded incorrectly as Adelaide Schomburgk 1867 ZMB 63847 Sydney Schomburgk 1868 ZMH R08213 514 Australia - 1861 ZMH R08212 763 Sydney Krefft Sep 1862 ZMUC R65270 Sydney - Sep 1862 ZMUC R65271 Sydney - ; Aug 1867 ZMUC R65272 Australia Don. Giinther 1920 ZSM 387/1920 NSW Destroyed in WWII 1928 ZSM 36/1928 NSW Destroyed in WWII Proc. Linn. Soc. N.S.W., 130, 2009 i SNe pati?’ Bray naa | ayleot ony arcane! 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Re a2 Melginen rsrby? ar czasy 54 Ohgtereatcred gantye (flan 2 “Tbe herent an oA ovkeat 1H Awkics the wav “WRITE ) ee 1 Trashed taal Aly 2 i alti” eeorec NR? a yin dbnipane ds A Te dj } Pri ag ex: hese 4, tage iit yd) Ties eal, Sia at, Gtacene Chin’ ROSK bree nie aides “ eT) Oonetien ed RYT Thaetdigais “« Sell irae: Chive a RII Wenn thing Calk Uhisewse Chow CHAS Bott Cok F. PPakgue Ce eal Wy ve tl Pond yader mach? a JETER i: Cagle ‘precy. ” pa ee Denis se Newel hon Mie 1 Sagi ne alle a Reconstructing Palorchestes (Marsupialia: Palorchestidae) - from Giant Kangaroo to Marsupial ‘Tapir’ B.S. MACKNESS School of Environmental Science and Management, Southern Cross University, PO Box 157, Lismore, New South Wales 2480, Australia. Mackness, B.S. (2008). Reconstructing Palorchestes (Marsupialia: Palorchestidae) - from Giant Kangaroo to Marsupial ‘Tapir’. Proceedings of the Linnean Society of New South Wales 130, 21-36. Since their initial description in 1873, palorchestid marsupials have been reconstructed in a variety of ways ranging from giant kangaroos, long-necked llama like-forms, bizarre okapians to their present popular image as quadrupedal marsupial ‘tapirs’. These reconstructions have resulted from an improved understanding of the phylogenetic position of Palorchestes, more complete fossil material and even the interpolation of supposed Australian Aboriginal renderings of these animals in Arnhem Land rock art. An examination of the timing of these different ‘views’ of Palorchestes has revealed that historical and social factors have also influenced how this animal has been visualized. Manuscript received 22 June 2007, accepted for publication 19 March 2008. KEYWORDS: history, Palorchestes, palorchestid, visual representation INTRODUCTION Attempts by vertebrate palaeontologists to reconstruct fossil animals are almost as old as the science that has informed such endeavours. In nineteenth century Europe, the French anatomist, Baron Georges Cuvier, gained a public reputation of being able to complete a “restoration from a single fossil fragment of complete skeletons of creatures long since extinct’ (Owen 1894:398). It appears, however, that Cuvier had only a marginal interest in attempting such reconstructions, dismissing them as too speculative (Coleman 1964, Outram 1984). Indeed, Cuvier didn’t publish any full reconstructions of prehistoric animals due primarily to his concern that such drawings would impact on his reputation as a scientist (Rudwick 1992). Across the channel, the so-called “British Cuvier’, Sir Richard Owen, earned similar accolades for his ability to reconstruct extinct animals from the most meager of remains. In one instance, Owen was said to have deduced the general form of the giant extinct New Zealand bird Dinornis from just “a six inch splint of bone with broken extremities” (Desmond 1975:101). Not all such palaeontological endeavours were so compelling however. When Cuvier was shown a tooth of the ornithischian dinosaur [guanodon, he identified the fossil as the upper incisor of a rhinoceros and later dismissed the metacarpal bones of the same animal as a species of hippopotamus (Delair and Sarjeant 1975). Owen’s work on Jguanodon was equally flawed. After being called on to supervise the sculpting of a life-size statue of the dinosaur, for the 1851 Great Exhibition of London, Owen not only posed the bipedal Jguanodon on all fours, but also placed its characteristic thumb spike on its nose (Desmond 1975). Although Cuvier was able to acknowledge his errors in identification before Mantell (1825) formally described /Jguanodon, Owen was not so fortunate. His anatomical faux pas were, and remain, highly visible thanks to the continued presence of the giant Iguanodon statue on its artificial island at Sydneham in London (Desmond 1975). In fact, almost a century and a half after its unveiling, Owen is still belittled over the anatomical inaccuracies of this reconstruction (Rudwick 1992) even though Owen was neither the first to reconstruct /guanodon nor the first to incorporate such maccuracies. Around 1835, for example, Mantell first visualized [guanodon as a type of a hypertrophied iguana (Williams 1991). Three years later, two further /gwanodon reconstructions were published in popular books on geology. George Nibbs completed a reconstruction as the frontispiece of George Richardson’s 1838 book, ‘Sketches in Prose and Verse’ while John Martin composed a gothic RECONSTRUCTING P4ALORCHESTES scene featuring three /gwanodon battling each other for Mantell’s, 1838 “Wonders of Geology’ (Rudwick 1992). Although significantly different from Mantell’s original iguana-like reconstruction, both followed his lead by picturing /gwanodon as a sprawling reptile with its thumb spike on its nose. While a paucity of fossil material has historically often been given as the reason for such errors in early reconstructions — in [guanodon’s case nothing more than a “few teeth and isolated bones” (Rudwick 1992:222) — other factors have also been implicated. At the time of Jguanodon’s discovery, the very concept of ‘dinosaur’ had not been formulated and the notion of extinct giant land reptiles was still novel (Delair and Sarjeant 1975:14). Further, given that there was also no demonstrated stratigraphic evidence that the Iguanodon fossils were anything olderthan Quaternary, it is perhaps not surprising that they were, at first, considered to be those of extinct mammals (Delair and Sarjeant 1975). Eventually, the existence of such giant land reptiles came to be accepted by scientists and even enshrined in the appellation Megal/osaurus or “great lizard’ — the formal name for the first of these creatures to be described (Buckland, 1824). As these giants had no living counterparts, they were understood using modern lizards as analogues and hence reconstructed as quadrupeds (Williams 1991). The first bipedal dinosaurs were not to be discovered for almost another two decades and on a different continent (Leidy 1858). As for the misplaced thumb spike, Mantell had originally indicated that the bone may be a dermal horn or tubercle but was convinced by unnamed authorities that the bone was a lesser horn of a rhinoceros (Delair and Sarjeant 1975). Even when Jguanodon was shown to be a giant reptile, it made more sense to place this ‘horn’ on the nose rather than on the hand given that there were no examples of similar thumb spikes in extant lizards. Desmond (1979, 1982), however, posits a deeper, political and perhaps even personal motives for Owen’s Crystal Palace reconstruction of /guanodon and the establishment of the taxonomic rank of Dinosauria (Owen (1841[1842]). This was to directly challenge the doctrine of Lamarckian transmutation, being espoused by many continental scientists and in England by his béte noir, Robert Grant of University College, London. Instead of giving the Crystal Palace statue the typical sprawling posture of all previous reconstructions, Owen stood his /guanodon erect like a mammal (Desmond 1982). By reconstructing it with such a modern stance, Owen hoped to discredit the doctrine of transmutation showing that present- day lizards and snakes represented a descent rather than an ascent as the ladder-like progression of the Dn, Lamarckian scheme demanded. Rupke (1994:133), however, contends that the establishment of the Dinosauria was nothing more than “the product of contemporary advances in taxonomic practices”. In Australia, the fossils of extinct giant marsupials, not dinosaurs, were the first to be studied and later reconstructed — primarily by overseas experts (Rich et al. 1985, Vickers-Rich and Archbold 1991). Among the earliest was Palorchestes, described by Owen (1873:387) as “the largest form of kangaroo hitherto found”. Its reconstructed skull was illustrated by Owen (1876) and then again in his seminal two volume work on Australian fossil mammals. In that work, Owen (1877) also provided a reconstruction of the country’s largest marsupial Diprotodon. As its feet were unknown at the time, the wily professor disguised these missing elements by hiding them in long grass. The foot bones were eventually found and described, almost a quarter of a century later, by Stirling and Zietz (1900). Modern reconstructions of Diprotodon differ little from the initial attempt by Owen except, of course, for the addition of the absent feet (Berganini 1964, Ruhen 1976, Quirk and Archer 1983). Other diprotodontid reconstructions have not been so readily accepted. The lack of recognizable postcranials of Zygomaturus meant that Gerard Krefft’s illustration of the animal, reproduced in Whitley (1966), was regarded as “curious speculation” by Archer (1984:677) while Lord and Scott’s (1924) reconstruction of the same animal was characterized as a “murky misconception” by Murray (1978:77), in spite of it being based on relatively complete fossil material (Scott 1915). The diprotodontoid Palorchestes, whilst being one of the first marsupials to be reconstructed, has also had the most varied reconstructions, being variously envisioned as a giant kangaroo (Owen 1876, Fletcher 1945); a gracile llama-like form (Bartholomai 1978); a bizarre okapian (Ford 1982); an elephantine- trunked quadruped (Flannery and Archer 1985); to its most recent guise as a marsupial ‘tapir’ (Quirk and Archer 1983) or ground-sloth-like creature (Long et al. 2003). Changes to how an animal has been reconstructed over time have normally been explained by reference to an increase in the availability of fossil material — “scientists of later periods have the benefit of more (and often better) specimens . . . than were available to their predecessors”(Rudwick 1992:220). The fossils of Palorchestes, however, are regarded as uncommon (Mackness 1995:606) or rare elements of fossil assemblages (Murray 1991:1106, Black 1997a:183), perhaps representing a solitary habit (Flannery 1983, Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS Flannery and Archer 1985, Black and Mackness 1999). The hypothesis that the extraordinary divergence in how Palorchestes has been reconstructed is due solely to changes in the amount of fossil material available has never been tested. Nor does such a suggestion allow for the influence of other factors even though these have been shown to have directly affected the visualization of other animals (Desmond 1979, Bakker 1988, Gould 1991, van Reybrouck 1998). This paper therefore seeks to systematically examine the major reconstructions of the marsupial ‘tapir’ Palorchestes, executed over the past 130 years, against the corresponding taxonomic understanding and available fossil material of the time in order to test the notion that changes in reconstructions of a particular animal result solely from improved fossil material and phylogenetic understanding and are independent of all other factor/s. The role played by palaeontological reconstructions in science communications is also discussed. MATERIALS AND METHODS Published reconstructions of Palorchestes from scientific and popular texts were digitally scanned and their main features rendered into line drawings. The taxonomic history of Palorchestes was chronologically arranged using summaries provided by Mahoney and Ride (1975) and Rich (1991). Details of fossils elements described were likewise listed in order of their publication following Woods (1958) and Rich et al. (1991), including those misidentifications that were used in the description of anatomical features of Palorchestes. Both these factors were compared against the line drawings of Palorchestes in order to ascertain whether there was any correlation between them. The possible effects of broader social and historical issues on each reconstruction were also considered. RESULTS Owen (1873) erected the genus Palorchestes on the basis of the anterior portion of a cranium, which included the rostrum. The holotype, collected by Dr Ludwig Becker from an unspecified deposit in Victoria, was named P. azael Owen, 1873. This locality has since been interpreted by Mahoney and Ride (1975) as the River Tambo in Gippsland. Owen assumed the animal was some sort of giant kangaroo as its cheek-teeth had longitudinal links between and in front of the transverse lophs (Archer 1984). These Proc. Linn. Soc. N.S.W., 130, 2009 features were later shown to have independently evolved in both palorchestids and kangaroos (Woods 1958). Nevertheless, Owen was convinced at the time that the new animal was a macropodid, a view reflected in his choice of its generic name, a conjunction of two Greek words which literally translate as ‘ancient leaper’ (Owen 1874:797). Two years later, Owen (1876) assigned further elements to P azael including a left and right mandibular rami, sacrum, caudal vertebra, innominate bone, femur, tibia, caleaneum and metatarsals, even though there was no field association with the holotype (Woods 1958). This same paper also contained the first published attempt to reconstruct Palorchestes in the form of an outline of its skull (Owen 1876, plate 20). The drawing (Fig. la), incorporated a realistic rendering of the holotype with a significant amount of the skull being inferred from extant kangaroos. This included the posterior portion of the cranium and the dentary. Surprisingly, although two mandibular fragments were assigned to Palorchestes in the same paper, they were not figured as part of the reconstruction but were used to justify the shape of the jaw as being most similar to Macropus, based on the changes in the depth of the fossil rami, rather than other extinct kangaroos such as Sthenurus and Protemnodon (Owen 1876). By reconstructing Palorchestes as a macropodid, Owen effectively obfuscated those features that would eventually come to be recognized as unique to palorchestids, such as the reduction of the nasals. Owen (1880a) described another species, P crassus from fluviatile deposits near Gowrie, south- east Queensland, on the basis of the symphyseal portion of a mandible with an anomalous condition in the molars of the right ramus. Lydekker (1887), however, found the condition absent in the left ramus and therefore synonomized P. crassus with P. azael. Woods (1958:182), in supporting Lydekker’s (1887) synonymy, further noted that the distortion originally described by Owen (1880a) was actually “postmortem fracturing, expansion and cementation with matrix”. A palorchestid palate from the Wellington Caves, New South Wales, named P rephaim by Ramsay (1885), was subsequently listed by both De Vis (1895) and Woods (1958) as P. azael. Consequently, the second valid palorchestid species to be described was P. parvus De Vis, 1895 from south-east Queensland. This new taxon appeared in De Vis’s (1895) paper on fossil macropodid jaws leaving no doubt that he shared Owen’s opinion that palorchestids were kangaroos. A premolar from Beaumaris Victoria identified by Hall and Pritchard (1897) as Palorchestes was later shown to belong to the Diprotodontidae (Stirton 1957). MB RECONSTRUCTING PALORCHESTES Figure 1. Historical reconstructions of Palorchestes from: a. Owen (1876); b. Fletcher (1945); c. Mur- ray (1978); d. Bartholomai (1978); e. Ford (1982); f. Quirk and Archer (1983). 24 Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS In 1912, the Trustees of the Australian Museum attempted the first three-dimensional reconstruction of Palorchestes using measurements from Owen and those from the mounted skins of living kangaroos (Fletcher 1945). The resulting sculpture stood almost three metres in height, even when posed in a resting position. Its imposing stature, when compared to that of living kangaroos, was said to have garnered much attention. This reconstruction was on display in the Museum for thirty-three years (Fletcher 1945). During the post-wars years, the higher classification of some mammal groups, including palorchestids, was reviewed by several workers. Simpson (1945) placed Palorchestes within the subfamily Macropodinae, following Owen’s lead, but the following year, Raven and Gregory (1946) moved it to the subfamily Sthenurinae. When Tate (1948) revised the kangaroos, he erected a new subfamily, the Palorchestinae, for Palorchestes. This meant that when the Australian Museum undertook a second supposedly more realistic reconstruction, taking into account “additional and important fossil remains” and to adopt “less misleading” assessments of how the animal should be modeled, Palorchestes was still thought of as a giant kangaroo (Fletcher 1945:363). The resultant model (Fig. 1b), was around 25% smaller than the 1912 original and photographed as the frontispiece of the Australian Museum Magazine (Fletcher 1945). Claims that this new museum model was the most accurate possible were somewhat tarnished however by errors in Fletcher’s (1945) accompanying text. He stated, for example, that Palorchestes was “first described in 1877 by Professor Sir Richard Owen, M.D., from the forepart of a crantum and portions of the jaw-bone with teeth” (Fletcher 1945:362-363) not in 1873 and based solely on a partial crantum as accepted by most other workers (Mahoney and Ride 1975, Mackness 1995, Black 1997a). Further, he interpreted the generic name Palorchestes to mean “the ancient dancer” (Fletcher 1945:362), even though Owen (1874a:797) specifically detailed its etymology. The greatest inaccuracies in the model, however, were to be exposed some thirteen years later. These were so significant that an embarrassed Australian Museum was forced to make a hasty and unceremonial disposal of their prized reconstruction (Archer 1984) with rumours still persisting that it is actually buried somewhere under Centennial Park in Sydney (M. Archer pers. comm.). The catalyst for the Museum’s precipitous action was a revision of Palorchestes by Woods (1958) who proposed that palorchestids were actually closer to diprotodontids than macropodids. The dentary of all Proc. Linn. Soc. N.S.W., 130, 2009 kangaroos possess a large mandibular foramen and masseteric canal. Both of these features were absent or suppressed in Palorchestes (Archer 1984). This meant that all the kangaroo-based reconstructions were incorrect and that palorchestids were most probably quadrupedal like other diprotodontids. Further, postcranials that had been attributed to Palorchestes in the past (e.g. Owen 1876, Gregory 1902, Scott 1916, Fletcher 1945) were shown by Woods (1958) to belong to either extinct kangaroos or wombats. The first undisputed palorchestid postcranial material was a series of caudal vertebrae of P. azael described by Bartholomai (1962), not in 1975 as claimed by Murray (1978). Five years after their description, a third palorchestid species, P painei Woodburne, 1967, was named from the Miocene Alcoota fauna of central Australia. Significantly, it showed the same extensive modifications to the rostral area that had been observed in P. azae/ and P. parvus by Woods (1958). In that same year, Stirton (1967) also formally recognized the Palorchestinae, which included Ngapakaldia and Pitikantia, as a subfamily within the Diprotodontidae. Archer and Bartholomai (1978) later raised this to familial status — the Palorchestidae. Further palorchestid postcranials were discovered in the seventies from a cave in the Wee Jasper area of New South Wales (Flannery and Archer 1985). These included a humerus and hindfoot which was subsequently prepared by the Australian Museum (Wells 1978). A humerus of P. azael was also reported from Victoria Cave, Naracoorte, South Australia by Wells (1975, 1978) along with phalanges and strange laterally-compressed scimitar-like claws, which Tedford of the American Museum of Natural History opined as being reminiscent of the extinct chalicotheres of the American Miocene. This led Wells (1978:109) to posit a tentative reconstruction of Palorchestes as “a large, quadrupedal grazing animal with longish limbs and plantigrade feet”. In the same year that Wells made his textual reconstruction, two new visual attempts were also published (Bartholomai 1978, Murray 1978). Both took account of Woods’s (1958) new phylogenetic understanding of palorchestids rejecting the earlier macropodid-based reconstructions. Murray’s (1978) sketch of a generalised Palorchestes (Fig. 1c), published in the specialist archaeological journal ‘The Artefact’, was based on the smaller Plio-Pleistocene palorchestid P. parvus. The reconstruction was part of a broader attempt to provide images of late Pleistocene fossil marsupials and a monotreme. Murray’s (1978, Fig. 12) sketch only included the head and shoulder D5 RECONSTRUCTING PALORCHESTES region, but a partial view of the entire animal was provided as part of a gallery of reconstructions (Murray 1978, Fig. 17). Following Woods’s (1958) re-description of P. parvus, Murray (1978:88) posited that Palorchestes would have had a “mobile upper lip indicated by the prominent pre-maxillary flange in the skull of P. parvus’’. It appears that Murray (1978:88) was also familiar with Fletcher’s (1945) article on the second model made by the Australian Museum as he repeated its error of interpreting the generic name of Palorchestes to mean ‘graceful dancer’. By contrast, Oakden’s scrapper board drawing of Palorchestes (Fig. 1d), for Bartholomai’s (1978) paper, was based primarily on the Miocene species P. painei. The catalyst for this reconstruction was the description of the cranium of P painei by Woodburne (1967); the preparation of further cranial material of the same species collected from the Waite formation during the 1974 Ray E. Lemley expedition of the Queensland Museum; and similar but less complete material of P azae/l and P. parvus held in the Queensland Museum (Bartholomai 1978:145). The reduction of the nasals, the elongation of the anterior of the palate and the presence of very large infraorbital foramina observed in these specimens led Bartholomai (1978) to postulate that all known species of Palorchestes probably had an extensive rhinarium or a tapir-like proboscis. Further, Bartholomai (1978) interpreted the narrow, deeply channeled mandibular symphysis as indicative of Palorchestes having had a long, flexible tongue. There were differences between the two reconstructions of Palorchestes, however, that could not be explained simply by the fact that they were based on different species. While Murray (1978:88) characterized Palorchestes as a ‘lightly buil[t] diprotodontid’, Bartholomai (1978) reconstruction was even more gracile with the longer neck making the animal look very llama-like. The position of the nares also differed, with those of Murray (1978) placed more posterior and superior to those in Bartholomai (1978). The latter was in line with Bartholomai’s (1978:148) assertion that Palorchestes may have possessed an “extensive rhinartum with anterodorsally directed nostrils”. Bartholomai’s (1978) Palorchestes was the first to feature a tapir-like trunk and also featured conspicuous vibrissae on the snout. By 1980, confirmation that the Wee Jasper material was indeed palorchestid came when a partial skeleton in the collection of the National Museum of Victoria was also shown to be that of Palorchestes (Flannery and Archer 1985). Although the Museum skeleton had no locality data, its association with some undisputed palorchestid teeth made the 26 specimen very important. Several of the bones in the skeleton had previously been labeled incorrectly by Scott (1916) as a giant species of wombat or wombat- like animal. Subsequently, other bones from Foul Air Cave at Buchan in eastern Victoria were also recognized as palorchestid. Given that the humerus of the Wee Jasper specimen was much smaller than the Buchan material, it was assumed that the Wee Jasper fossils represented P parvus while the Buchan bones were those of the larger P. azae/ (Flannery and Archer 1985). The identification of this additional postcranial material enabled a full reconstruction of Palorchestes as a quadruped. In 1981, Stahel produced a stipple drawing of an entire animal for an article published in a University newsletter (Archer 1981). This illustration was used the following year as the basis of a reconstruction (Fig. le) by Ibraham for an article in the science magazine ‘Omega Science Digest’ titled ‘The strange creatures of ancient Australia’ (Ford 1982). What is significant about both drawings is that they embodied a rather “chimeraesque’ understanding of Palorchestes, demonstrating a concomitant “high coefficient of weirdity” (Archer 1984:670). The overall body outline was rather ‘okapian’ with the hind-quarters lower than the front and the neck long and giraffid-like. The ‘bizarre herbivorous animal’ was Said to be as “large as a horse . . . [with] a trunk- like structure on its face .. . kangaroo-like teeth . . . [a] long giraffe-like tongue and . . . phenomenally huge sharp claws” (Ford 1982:84-85). These sharp koala-like claws were even thought, for a brief time, to represent an adaptation to climbing in trees like modern-day sloths but the idea was rejected when the huge size of Palorchestes became apparent (Archer 1984:670). These speculative views of Palorchestes were informed by palaeontologist Mike Archer who, just one year later, was involved in the production of another reconstruction that directly challenged many of the assumptions inherent in the ‘okapian’ model (Archer 1984). The rethink of how Palorchestes should be reconstructed was prompted by several factors including the identification of additional fossil elements and the opportunity to further refine or challenge aspects of previous reconstructions. The neck length of the Stahel and Ibraham reconstructions, for example, was deemed too long after the discovery that palorchestid cervical vertebrae were not elongate like that of giraffids (Archer 1984:670). Likewise, the size of the trunk was also thought to be over-inflated and consequently reduced with the tail likewise being shortened. These changes were encapsulated in a new rendering of Palorchestes which Archer (1984:670) Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS judged to be the “best” to date, acknowledging however that his opinion was biased, given his involvement in its formulation. The reconstruction, executed by Schouten (Fig. 1f), appeared in a book on prehistoric animals published by the Australian Museum (Quirk and Archer 1983). Schouten presented a composite view of the head and front feet of P. azae/ along with a full-view of the animal ripping bark from a tree. Beneath this illustration, a further sketch was provided to demonstrate how Palorchestes may have used its tongue to strip vegetation off branches. The body shape of Schouten’s Palorchestes was much more diprotodontid-like and its size more like that of a bull. The reconstruction also highlighted Palorchestes’s massive forearms; its rapier-like claws and tapir-like trunk. The text accompanying the new reconstruction was titled “unique trunked giant” and contained the first explicit connection between Palorchestes and Aboriginal people. Flannery (1983:54), who penned the text, suggested that Palorchestes may have been the inspiration behind the legend of the bunyip and that newly arrived Aboriginals may have had second thoughts about settling after seeing one of these giant marsupials. Further, Flannery (1983:54) claimed that Aboriginal people and Palorchestes had “co-existed in Australia between about 40 000-20 000 years ago”. In 1984, three different reconstructions of Palorchestes were executed by Murray, but in very different contexts. The first was a drawing of a generalized palorchestid (Fig. 2a) as part of a family tree of diprotodontoids presented in a children’s book ‘Australia’s prehistoric animals’ (Murray 1984a). Both Palorchestes and the mid-Miocene Neapakaldia were shown on the same blue branch representing the Palorchestidae (Murray 1984a). In contrast to his 1978 reconstruction of Palorchestes (Fig. 1c), however, Murray’s new depiction had a much longer tapir-like trunk. This interpretation was” justified with the inclusion of a diagram showing the similarities between the skull and trunk of a tapir and that suggested for Palorchestes. Murray’s illustration differed from Schouten’s (Fig. 1f) in having a longer tail but smaller body. Murray was also the first to explicitly use the term “tapir-like marsupial” (Murray 1984a:20). Murray’s second reconstruction was specifically of P. azael (Fig. 2b) and was published in a book on Quaternary extinctions. As with Ford’s (1982) characterization, Palorchestes was once again presented as a composite animal only this time it was said to have “tapir, chalichothere, pantodont and sloth-like features” (Murray 1984b:608). The “large kangaroo-like tail” of P azael was highlighted, citing Proc. Linn. Soc. N.S.W., 130, 2009 Bartholomai (1962) and a personal communication from the same author, while Archer and Bartholomai (1978) were quoted as the source of P. azae/ being “equipped with huge, curved, laterally compressed claws” (Murray 1984b:608). The overall body size of Murray’s P. azael was much more massive than his more generalized drawing (Fig. 2a) and featured a long flexible tongue. Fossil remains of P. azael were regarded by Murray (1984b) as not especially common but widely distributed, with specimens of P azael from Pulbeena Swamp in Tasmania, (54 200+11 000 - 4 500 yr BP) listed as a recent occurrence of the taxon (Banks et al. 1976). Flannery’s (1983) suggestion that Palorchestes and Aboriginal people lived contemporaneously was seemingly validated in 1984 when a large Aboriginal painting (Fig. 2c) was tentatively identified asa possible representation of the extinct marsupial (Murray and Chaloupka 1984). The painting, discovered in Deaf Adder Gorge, Arnhem Land in 1976, was part of a tradition called the Large Naturalistic Animal Style (sensu Chaloupka 1993), which included depictions of animals now extinct from the Australian mainland such as thylacines and Tasmanian devils (Calaby and Lewis 1977, Lewis 1977, Clegg 1978). Some of the features used by Murray and Chaloupka (1984) to identify the painting as Palorchestes included: 1) the considerable attention given to the tongue including small lines which were said to perhaps represent items of food such as leaves or insects; 2) the detail given to the claws and the angled calcaneal joint; and 3) a lack of ears. Two anomalous breast-like projections under the body were explained as “stylised attempts to show a long shoulder mane or shaggy long hair” (Murray and Chaloupka 1984:114). A smaller animal besides the larger painting was said to represent a joey of the extinct marsupial. Murray and Chaloupka (1984) compared the Palorchestes painting with those of introduced animals such as those found previously in Cape York (Trezise 1971) as well as a variety of megafaunal species. In suggesting that the painting represented a Palorchestes, Murray and Chaloupka (1984:115) were extremely circumspect however, stating that “maybe it [the painting] represents Palorchestes” but “it must be made very clear that the connection at present is of the most tenuous kind”. They even suggested that “there may not be much gained by attempting to compare this unique and intriguing painting with perhaps the most poorly known species in the megafaunal assemblages” (Murray and Chaloupka 1984:112). In spite of such tentativeness, however, and in spite of a serious challenge to both the methodology and assumptions used (Lewis 2a RECONSTRUCTING PALORCHESTES Figure 2. Further reconstructions of Palorchestes from: a. Murray 1984a; b. Murray 1984b; c. Arnhem Land ‘Palorchestes’ from Murray and Chaloupka (1984); d. Murray and Chaloupka (1984); e. Rich et al. (1985), f. Long et al. (2003). 28 Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS 1986, Mackness, unpublsihed data), the painting has been promoted as a credible example of megafauna depiction by Aboriginal artists (Chaloupka 1993, Flood 1997). A third Palorchestes reconstruction (Fig. 2d) by Murray appeared in his joint paper with Chaloupka on rock art. What was unique about the reconstruction was that certain features were specifically added to match the supposed Aboriginal representation of Palorchestes. The most obvious of these was a mane of long hair protruding below the line of the abdomen to match the anomalous projections of the painting (Murray and Chaloupka 1984). This feature was not present in any of Murray’s previous 1984 reconstructions. The ears were also placed so that they didn’t project beyond the outline of the head to likewise match the painting. In Murray’s generalised Palorchestes (Fig. 1a), the line of the ears was clearly shown projecting above the head. In support of such modifications, the authors restated Clegg’s (1981:313) assertion that “if a well executed drawing of potentially great antiquity best matches a good restoration of an extinct species , then that may well have been the target species”. While invoking this “Occam’s Razor of rock art analysis” as justification for their identification of a Thylacoleo drawing, Murray and Chaloupka (1984:115) regarded the evidence for the Palorchestes drawing as being “less satisfactory” however. While the reconstructions of Palorchestes by both Schouten and Murray featured relatively short tapir-like trunks and diprotodontid-like bodies, Knight’s (Fig. 2e) composite illustration of P. azael and P. parvus, published in Rich et al. (1985), featured much longer trunks, body shapes more reminiscent of myrmecophagids and rhinoceros- like tails. Knight actually completed the illustration in 1982, around the same time that the Stahel and Ibraham reconstructions were published. The text accompanying the illustration, by Flannery and Archer (1985), provided the first detailed description of palorchestid postcranials along with a sketch of the articulated arm bones and a rear view of the humerus. Flannery and Archer (1985) argued that the front legs of palorchestids were unusual, relative to other marsupials, because of a greatly enlarged area for the attachment of the pectoralis muscle which formed a high, hooked process. The ulna of both species was said to be almost solid with only a tiny marrow cavity. The nature of the articulation between the lower and upper arm bones in P. azael was such that it appeared to indicate an immobile elbow with the front legs being permanently locked in a partly flexed Proc. Linn. Soc. N.S.W., 130, 2009 position, strengthening the already massive forearms. The smaller P. parvus, however, appeared to have a slightly more flexibility in this joint. The authors also drew attention to the highly mobile fingers that each bore a massive, sharp, laterally-compressed claw similar to that of a koala but far larger. Flannery and Archer (1985) interpreted these claws as suitable for ripping, tearing or climbing but not for digging. By comparison, the authors considered the hindlimb of Palorchestes to be far less robust. The fourth and fifth toes were equipped with the same kind of massive claws seen on the fingers of the hands but toes two and three were reduced in size and syndactylous, perhaps used for grooming. Flannery and Archer (1985) also suggested that Palorchestes may have possessed a clawless opposable great toe similar to that seen in possums. Overall they suggested that Palorchestes filled a niche similar to that of elephants or the extinct ground sloths of the Americas, using its narrow and elongate tongue in conjunction with its trunk, to strip leaves off trees and bushes. Once again, an explicit connection was made between Palorchestes and Aboriginal people with the suggestion that the “exceptionally powerful forearms, massive claws and bizarre head would surely have been enough to have inspired the legend of the bunyip — or at least a few nightmares among Australia’s first Aboriginal inhabitants” (Flannery and Archer 1985:236). The composition of the Palorchestidae was challenged by Murray the following year with the description of the lamb-sized palorchestid Propalorchestes from mid-Miocene deposits of Bullock Creek Local Fauna, Northern Territory and several Oligo-Miocene sites at Riversleigh, Queensland. Doubts had previously been cast by Archer and Bartholomai (1978) and Archer (1984) about the monophyly of the Palorchestidae. Aplin and Archer (1987), in their review of marsupial systematics, had placed palorchestids in their present position within the Vombatiformes. A further reconstruction of Palorchestes (Fig. 3) was executed by James Reece for a popular book on prehistoric life by Mackness (1987). Reece combined the reconstructions of Schouten and Knight to produce a hybrid image that adhered to a by now standard formula for illustrating Palorchestes with a diprotodontid body, sharp claws and tapir-like trunk. Such visual codification, called conventionalization by Rudwick (1992) enabled those viewing the animal to instantly recognize it as Palorchestes. In 1990, Murray described another species of Propalorchestes and concluded that members of that genus were the plesiomorphic sister-taxon of 2 RECONSTRUCTING PALORCHESTES i) / Lt if i} y Pigia } i | 34 ae ‘ ~ \ V4 Les | | Ga a | | se } al | | / Ws I | / Af i ‘i / aa / | | > - att 2 | | S veg f % we / | fy be y} { fi # s oe L , | Ne @ i. 17 = ety | ol & | on | N ~~ | | { i i | | =, he r /) | tI | | \ , | i | { | on | oe \ | | | \ Hi | aa , : | ; \ get Ky y aN \ ¥ “a ae i 3 ‘a 1 _ | ; teen =| == i / ro ) / | SA, N \ ule ; | | et ee aie (eee a eee i ra } | ves \ LG ' | = ga! ee oe | | sine Saar | < ‘SS WN IN | Figure 3. Reconstructions of Palorchestes from Mackness (1987). Palorchestes while Ngapakaldia and Pitikantia should be regarded as primitive members of the Diprotodontidae (Black 1997a). Five years later, a new species of palorchestid, Palorchestes selestiae, was described from the early Pliocene Bluff Downs Local Fauna on the basis on an isolated M! (Mackness, 1995) with a fifth species, P anulus described just two years later by Black (1997a) from the early-late Miocene Encore Local Fauna, Riversleigh, again on the basis of an isolated M'. The most recently described palorchestid, P. pickeringi, was recovered by Piper (1996) from Pliocene and early Pleistocene deposits of Victoria. It is represented by a significant 30 amount of fossil material and has also possibly been identified from Queensland (Hocknull et al. 2007). By the last decade of the twentieth century, the term “marsupial tapir” had become firmly entrenched as the popular name for palorchestids (Murray 1991) even though alternative descriptors such as “marsupial tree-fellers” had been proposed (Flannery 1994). The visual codification of Palorchestes reconstructions continued to be refined with the most recent reconstruction of P. azael (Fig. 2f), executed by Anne Musser and published in Long et al. (2003), perhaps being the apogee of how the animal should be depicted. Musser’s illustration did not show an Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS exaggeratedly long tongue or a trunk capable of being bent back on itself as illustrated by Schouten. The forearms were shown to be immobile following Flannery and Archer (1985), while the tail was more like that proposed by Murray (1984b). The explicit connection between Palorchestes and the eutherian Tapirus was also being down-played with extinct ground sloths now being the dominant analogue. This suggestion, first raised by Archer (1984) and Murray (1991), was visually encoded by the depiction of Palorchestes walking on the sides of its feet or on its knuckles. Long et al. (2003) also included an illustration of the skull of P painei showing its fragmentary nature, linking the real with the inferred in a similar manner to that first employed in Owen’s (1876) first reconstruction almost a hundred and thirty years previously. DISCUSSION The veracity of palaeontological reconstruction is underpinned by a specific methodology which is supposedly deployed with each attempt to illustrate a prehistoric creature. Murray (1978:77) characterizes “serious” reconstructions as only those that are based on “detailed anatomical build up of soft tissues”. This requirement challenges most reconstructions as very few conform to such rigor. Schouten visualized this same process using Diprotodon as an example in Quirk and Archer (1983). It should be noted, however, that it would have been singly impossible for any one artist to have the detailed anatomical knowledge required to undertake similar soft tissue build ups of all the other animals illustrated in that work. Rudwick (1992:221) provided yet another outline of the methodology suggesting it occurs in the following sequence:- 1) the selection of suitable fossil bones for assembly of a partial skeleton of a particular individual; 2) the reconstruction ofa complete skeleton representative of the species, based generally on the remains of many individuals; 3) reconstruction of a generalised complete individual body with inferences about the animal’s unpreserved muscles and other soft parts, based partly on anatomical analogy with related living forms; 4) and finally inferences about the animal’s dynamic mode of life and habits, based partly on functional analysis of its anatomy and on physiological analogy with related living forms. Rudwick (1992:221) posits that the outcome of such a sequence is “a cascade of representations that are progressively bolder—yet still well-founded— reconstructions of the unobservable prehuman past Proc. Linn. Soc. N.S.W., 130, 2009 . . . progressing from the observed to the inferred, from the specific and contingent to the general and idealized”. Changes in successive attempts to portray the same animal are simply “attributed to the discovery of more and better specimens that are relevant to that reconstruction” (Rudwick 1992:220). Latour (1986:17), however, from whom Rudwick (1992) derived the notion of “cascade”, uses the term in a much different sense. For Latour (1986:17), the sequence of reconstructing a prehistoric animal results in a “cascade of ever simplified inscriptions [visual representations] that allow harder facts to be produced”. Therefore, it is the selection of bones from a collection to be used in the description of a new species or the reconstruction of a complete skeleton from bones held in several museums over a wide geographic locality that allow scientists to make “bolder” reconstructions. When a pile of individual elements are coalesced into a published type description or into an articulated form, they became a single entity of “the type of . . .” or “the skeleton of . . .” with all its associated eidetic qualities. This process of accumulation and simplification is only useful however when there is confidence that the meaning of each coalescence has been stabilized (Pinch 1985). If it hasn’t, then all subsequent layers that are built upon it risk collapsing like a veritable ‘house of cards’ should the underlying assumptions prove to be unstable or incorrect. Such was the case with Owen’s (1876, 1877) reconstruction of Palorchestes as a macropodid. While in hindsight, it may seem that Owen made a grave error in his classification of the animal, Fyfe and Law (1988:1) caution that“... both the processes that lead to the creation of depictions, and the way in which they are subsequently used, have to be studied in their historical specificity”. With Palorchestes, several factors mitigated against Owen recognizing its ‘true’ taxonomic affinities. The partial cranium used as the holotype, for example, lacked those features, such as the reduction and retraction of the nasals, which would eventually be regarded as autapomorphies for palorchestines. Indeed, it wasn’t until almost a century later, after Woods (1958) had revised the genus and Woodburne (1967) had described P. painei, that suitable material became available to elucidate such characters. The presence of longitudinal links between and in front of the transverse lophs, while used by Owen (1874) to justify Palorchestes as a kangaroo, has since been shown to be convergent with at least two zygomaturine genera — Maokopia Flannery, 1992 and a new, as yet unnamed, Plio-Pleistocene species from eastern Australia (Black and Mackness 31 RECONSTRUCTING PALORCHESTES 1999. Mackness, unpublished data) possessing similar links. Flannery (1992:325) postulates that the development of “anteroposteriorly directed linking is an adaptation to a more abrasive diet’. Similarly, it wasn’t until the early part of the twentieth century that Abbie (1939) demonstrated that the presence of the masseteric fossa was a feature that united all macropodids. The fossil rami described by Owen (1876) lacked this relevant portion. Archer (1984) rightly concluded that the absence of such a feature in palorchestids didn’t preclude the possibility that they were still a plesiomorphic sister group of kangaroos. It wasn’t until Murray’s (1986, 1990) description of Propalorchestes and detailed biostratigraphical research into the Riversleigh Local Faunas by Black (1997b) that the taxonomy of palorchestids obtained some sort of stability with many authors (e.g. Archer and Bartholomai 1978, Archer 1984, Murray 1990, Mackness 1995) having previously cast doubt about the phylogenetic make-up of the group. The first major rethink about how Palorchestes should be reconstructed was not so much a result of additional and better fossil evidence becoming available as required by Rudwick’s (1992) sequence, but rather a reassessment of existing museum material and a consequential re-interpretation of its phylogenetic affinities (Woods 1958). This conforms to Latour’s (1986) notion of a ‘cascade’ with Wood’s (1958) coalescence providing a stable platform for harder facts to be produced. When new fossil material was collected by Woodburne (1967) and Bartholomai (1978), it was therefore added to the already stable platform of ‘palorchestids as diprotodontoids’. In particular, Bartholomai’s (1978) interpretation that the rostral area of palorchestids may have supported a tapir-like proboscis or extensive rhinarium provided the basis for the interpretation of palorchestids as marsupial ‘tapirs’. The lack of unequivocal palorchestid postcranials, however, apart from those described by Bartholomai (1962), meant that only the head region was known well enough for Bartholomai (1978) and Murray (1978) to attempt reconstructions — except for one very generalized body view (Murray’s 1978, Fig. 17). Even after palorchestid postcranials had been discovered and identified from caves in New South Wales, Victoria and South Australia in the 1970’s, their lack of publication meant they were effectively unavailable for use in reconstructions except for those few who had access to the relevant museum collections and the detailed anatomical knowledge to interpret what individual elements represented. To this day, the only description of these fossils is the popular account by Flannery and Archer (1985) in Rich et al. (1995). Sy The temporal lag of almost a decade between the discovery of these fossils and their incorporation into reconstructions also suggests that the relationship proposed by Rudwick (1992) may not be as straight forward as first thought. While some delay is to be expected, to allow for the preparation, study and publication of fossils, the postcranials of Palorchestes were never published in a peer-reviewed journal. Further, the most diverse representations of Palorchestes occurred between 1981 and 1983 (acknowledging that Knight’s reconstruction was completed in 1982) after the concept of palorchestids as diprotodontoids was stabilized by Woods (1958). The various attempts at reconstruction may, in part, be due to scientists using them as heuristic devices to test various anatomical options. The fact that palaeontologists Archer and Flannery, supervised all these divergent ‘views’ of Palorchestes perhaps bears this out. Van Reybrouck (1998), in his study of Neanderthal reconstructions, suggests that the intellectual zeitgeist may also affect how an organism is visualized. The publication of the various reconstructions of Palorchestes coincided with what Tedford (1991:76) characterizes as the “coming of age” of Australian vertebrate palaeontology with many academic institutions launching indigenous study programs at that time. Concomitantly, it was also a time when attempts were being made to raise the profile of the discipline in order to attract new students to the nascent palaeontological programs being offered at Universities (Vickers-Rich and Archbold 1991, Tedford 1991); to raise funds for research; and to mobilize and educate the general public (Quirk and Archer 1983, Rich et al. 1985, Mackness 1987). Perhaps not surprisingly, these popular texts featured creatures with superlative values such as the oldest, the largest or in Palorchestes’s case, the weirdest (Archer 1984:670). Part of the reason Palorchestes came to be reconstructed in so many guises was its ‘weirdness’ when compared to other marsupials. As well as being co-opted as a ‘poster child’ to demonstrate the uniqueness of Australia’s past, Palorchestes was included in some seminal debates about Aboriginality concerning the interrelated topics of land rights, environmental management and the extinction of the megafauna. Questions about the antiquity of Aboriginal settlement of the Australian continent had followed the widespread availability of radiocarbon dates (Mulvaney and Kamminga 1999) and in particular the dating of the Lake Mungo burials. A date of more than 40 000 years became a “slogan for indigenous people” (Gillespie 2004:1) and mobilized in legal arguments about rights to land (Yunupingu Proc. Linn. Soc. N.S.W., 130, 2009 B.S. MACKNESS 1997). The contemporaneity of Aboriginal people and extinct megafauna was another plank in this argument with suggestions that Palorchestes was the subject of the bunyip legend (Flannery 1983, Flannery and Archer 1985) and its supposed representations in rock art (Murray and Chaloupka 1984) adding credence to such claims. While Owen (1880b) was amongst the first to implicate Aboriginal people and the extirpation of the Australian megafauna, the early eighties saw the emergence of a full blown debate on the issue (Horton 1979, 1980; Martin and Klein 1984), a subject that continues to provoke controversy two decades later (Flannery 1994, Horton 2000, Roberts et al. 2001, Wroe et al. 2004). Consequently, while fossil discoveries and reinterpretations of phylogenetic relationships have played an important part in the varied reconstructions of Palorchestes, other broader factors have also been implicated. No matter what these influences are, however, they only become relevant if a particular reconstruction continues to be deployed. Corrigan (1988) contends that every time someone reproduces a reconstruction it becomes imbued with power. The context of reproduction can also play an important part in how a reconstruction is judged. Schouten’s 1983 reconstruction of Palorchestes azael has, until recently, held sway not only because it supposedly best matched the fossil evidence and was the most sophisticated rendition (Archer 1984) but also because it appeared in a book published under the imprimatur of the Australian Museum, one of the nations leading scientific institutions. The most recent reconstruction by Musser in Long et al. (2003) has yet to gain the same widespread exposure of Schouten’s effort but it obviously has only been in circulation for a short time. Its eventual hegemony also rests on the acceptance of the ground sloth analogy, explicit in the reconstruction rather than the existing and long-standing marsupial ‘tapir’ model. Latour (1987:258) suggests that *. . to determine the objectivity or subjectivity of a claim [like that made by a scientific illustration] . . . we look not for their intrinsic qualities but all the transformations they undergo later in the hands of others’. Consequently, future reconstructions of Palorchestes will not just be judged by whether or not they best fit the palaeontological information available but also whether they are reproduced in wide enough contexts to be accepted. ACKNOWLEDGMENTS The author wishes to thank Bill Boyd, Southern Cross University; Sue Hand, University of New South Wales and Proc. Linn. Soc. N.S.W., 130, 2009 Errol Vieth, Central Queensland University for providing helpful comments on the manuscript. Greg Luker, Southern Cross University, undertook the line reproduction of the different Palorchestes reconstructions. Katarzyna Piper provided valuable access to her work on palorchestid marsupials. Glenda Kemmis, Southern Cross University Library sourced the many reference articles used in the paper. The study was supported, in part, by a postgraduate grant to the author from the School of Environmental Science and Management, Southern Cross University. REFERENCES Abbie, A.A. (1939). A masticatory adaptation peculiar to some diprotodont marsupials. Proceedings of the Zoological Society of London Series B. 109, 261-279. Aplin, K. and Archer, M. (1987). Recent advances in marsupial systematics with a new syncretic classification. In ‘Possums and Opposums: Studies in evolution’ (Ed M. Archer) pp. xv-lxxii. (Surrey Beatty and Sons and the Royal Zoological Society of New South Wales: Sydney). Archer, M. (1981). Hunting ancestors in the possum dreamtime. 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A basal actinopterygian fish from the Middle Devonian Bunga Beds of New South Wales, Australia. Proceedings of the Linnean Society of New South Wales 130, 37-46. A partial articulated skeleton of a basal actinopterygian fish is described from the Middle Devonian Bunga Beds of New South Wales. The specimen represents a new species and is questionably assigned as a congener of Howqualepis rostridens from the Middle Devonian of central Victoria. This represents the first record of an articulated postreranium of a Devonian ray-finned fish from New South Wales. The pectoral fin of Howqualepis is also redescribed based on a re-examination of Victorian material. The fin is broader in shape and less extensively unsegmented than previously recognised. The close similarity of the new form with contemporaneous taxa from Victoria and the Aztec Siltstone of Antarctica adds to an already wide body of evidence supporting a regionally endemic freshwater vertebrate fauna in the Middle Devonian of Eastern Gondwana. Manuscript received 7 Feb 2008, accepted for publication 22 October 2008. KEYWORDS = Actinopterygians, Bunga Beds, Devonian, fish, Howqualepis, New South Wales INTRODUCTION In stark contrast to their modern abundance and diversity, actinopterygians are a sparse component of most Devonian vertebrate assemblages. Australia is notable in producing some of the finest fossils of Devonian actinopterygians, the best known of which are exceptionally preserved specimens from the Frasnian Gogo Formation of northern Western Australia. Included within the assemblage are Moythomasia durgaringa (Gardiner & Bartram 1977, Gardiner 1984), the currently preoccupied “Mimia” toombsi (ibid), Gogosardina coatesi (Choo et. al, in press) and at least two additional forms (Choo, in prep). Southeastern Australian fossil sites have also produced a substantial amount of early ray-finned fishes. The first record of Australian Devonian actinopterygians consisted of the isolated scales of Ligulalepis toombsi from the Lower Devonian Taemas-Wee Jasper Limestones of New South Wales (Schultze 1968). A subsequently discovered braincase and skull-roof was assigned to this genus (Basden et al. 2000, Basden & Young 2001). Long (1988) described Howqualepis rostridens based on numerous specimens from the Givetian Mt Howitt fauna of central Victoria (age revised in Young, 1999). Adding to this Eastern Australian record is an incomplete but articulated fossil that was recently discovered by Gavin Young from the Middle Devonian Bunga Beds, near the shoreline at Bunga Beach in south coastal New South Wales. This represents the first discovery of an articulated Devonian actinopterygian postcrantum from New South Wales. Subsequent repeated searches failed to recover additional material of this form (Gavin Young, pers. com.). GEOLOGICAL SETTING The Bunga Beds represent a thinly bedded sequence of carbonaceous shale and sandstone that comprises the lowest section of an extensively fossiliferous Devonian sequence (Fergusson et al. 1979, Young 2007). Young (2007, figs 1, 2) provides and up to date account of the lithology, fossil assemblage and possible age of the Bunga Beds. The age of the unit is poorly constrained and probably older than the Late Devonian age stated in recent A DEVONIAN ACTINOPTERYGIAN FISH literature (Cas et al. 2000, Giordano and Cas 2001, Rickard and Love 2000). The dark shales of the Bunga Beds are highly fossiliferous with abundant plant material and uncommon vertebrate remains (Young 2007, fig. 3), possibly representing a deepwater lacustrine depositionalenvironment. The fossilfish fauna includes ischnacanthid acanthodians (Burrow 1996), several taxa of chondrichthyans including Antarctilamna prisca (Young 1982), originally described from the Givetian Aztec Silstone of Antarctica, and a possible tetrapodomorph sarcopterygian (Young 2007, table 1). The fossil ichthyofauna of the Bunga Beds seems impoverished due to the apparent absence of placoderms and dipnoans that are abundant in other southeastern Australian sites of a similar age. MATERIALS AND METHODS The fossil was recovered as a natural mould set within a matrix of dark shale. After collection, the Specimen was split into part and counterpart and bone remnants removed. Bone margins were exposed with mechanical preparation and the impressions of the fish were examined using a latex rubber cast whitened with ammonium chloride. For comparison, fresh latex casts were made from the following specimens of Howqualepis rostridens in Museum Victoria (MV) = P.160745A, P.160782A, P.160788, P.160792B, P.160811, P.160822A, P.160851B, P.160857. Abbreviations for actinpterygian dermal bones and other structures used in the text and figures are as follows: an.f, anal fin; Br. 1, 1st branchiostegal ray; Br. 7, 7th branchiostegal ray; c.ful, caudal (basal) fulcra; Clav, clavicle; Clth, cleithrum; cw.lep, cutwater of short lepidotrichial segments; d.lep, probable dorsal lepidotrichia; f.ful, fringing fulcra; nm, notochordal mass of caudal fin; Op, operculum; Sop, suboperculum; pec.f, pectoral fin; pel.f, pelvic fin; pseg, segmented posterior lepidotrichia on pectoral fin; tfr, terminal fringe of fine branching segments on pectoral fin; vhl, ventral hypochordal lobe of caudal fin; useg. unsegmented proximal lepidotrichia on pectoral fin. SYSTEMATIC PALAEONTOLOGY CLASS OSTEICHTHYES Huxley, 1880 SUBCLASS ACTINOPTERYGII, Cope, 1887 Family Howqualepididae Long, Choo and Young, 2008 38 Diagnosis (revised) Basal actinopterygians with an open spiracular slit bordered by the intertemporal, dermosphenotic and supratemporal. Intertemporal is very small (less than 1/3 the size of parietals). Pineal foramen present on anterior half of the median frontal contact. Dermosphenotic is elongate and tripartite. Suboperculum has a prominent anterodorsal process. Body form is elongate and fusiform. Squamation macromeric; scales are rhombic with linear ganoine ornamentation. Fringing fulcra are spine-like terminal sections of the anterior fin rays, lacking median contact between the hemilepidotrichia. Longest anterior pectoral fin rays are proximally unsegmented for over 60% of their length. Median scute series on dorsal and ventral surface do not extend anteriorly to reach the head. Remarks Diagnosis slightly modified from Long et. al (2008) to incorporate the revised description of the pectoral fin and fringing fulcra of Howqualepis presented below. Genus ?Howqualepis Long, 1988 ? Howqualepis youngorum sp. nov. Etymology After Professor Gavin Young (ANU) who discovered the holotype specimen and Mr Ben Young for conducting both the preparatory work as well as the key photography of the specimen. Repository The type and only known specimen is lodged in the collections of the Department of Earth & Marine Sciences, Australian National University, Canberra, represented in the text by the prefix ANU V. Holotype. ANU V2929a, b, an incomplete, partially articulated fish preserved laterally in part and counterpart. Consists of an incomplete opercular- gular series, cleithrum, clavicle, scales and all fins except the dorsal fin (Figs. 1-4). Collected by Gavin Young (ANU) from the Bunga Beds at Bunga Beach, south of Bermagui, New South Wales. Diagnosis A Howqualepis with more than 54 primary lepidotrichia on the anal fin and porous ornamentation on the cleithrum and clavicle. Proc. Linn. Soc. N.S.W., 130, 2009 B. CHOO Remarks Tentatively assigned to the genus Howqualepis. The extensive unsegmented pectoral lepidotrichia of ?Howqualepis youngorum sp.nov separates this taxon from all other Devonian actinopterygians except Howqualepis rostridens Long, 1988, Donnrosenia schaefferi Long, Choo and Young, 2008, and Tegeolepis clarki Newberry, 1888. ?H. youngorum differs from Donnrosenia in that the unsegmented fin-rays account for more than 75% of the total length of the pectoral fin. ?H. youngorum differs from Tegeolepis in possessing macromeric squamation, long-based pelvic fins and a segmented terminal fringe on the pectoral fin. Separable from H. rostridens in having porous (as opposed to entirely linear) ornament on the pectoral girdle and in having a larger anal fin (54+ vs 45 primary lepidotrichia). DESCRIPTION Overall body form ANU V2929 is preserved in lateral aspect (Fig. 1). The anterior part of the specimen terminates at an oblique breakage margin, with elements of the opercular-gular series and pectoral girdle preserved along with the pectoral fin (Fig. 2). 2.5 cm behind this is an incomplete pelvic fin with patches of squamation present above and to the rear of the fin (Fig. 3). The largest preserved segment comprises the rear section of the fish, including well preserved anal and caudal fins along with extensive squamation (Fig. 4). The preserved sections suggest a highly elongate, fusiform body form similar to that of Howqualepis rostridens (Long 1988) and quite unlike the more compact and robust form of “Mimia” or Moythomasia (Jessen 1968, Gardiner 1984). As preserved, the fossil measures slightly less than 12 cm from the anterior preserved edge of the clavicle to the posteriormost caudal scales. Assuming that the missing portions of the fish were of similarly proportions to that of Howqualepis rostridens, the complete fish would have measured about 14 cm from snout to caudal peduncle. Opercular-gular series A section of the dermal operculo-gular series of ANU V2929 is preserved in articulation and comprises Figure 1. ?Howqualepis youngorum sp. nov. a. photograph and b. line drawing of holotype (ANU V2929A) showing the entire preserved fossil in lateral view. The specimen is a latex cast whitened with ammonium chloride. Proc. Linn. Soc. N.S.W., 130, 2009 39 A DEVONIAN ACTINOPTERYGIAN FISH Figure 2. ?Howqualepis youngorum sp. nov. a. photograph and b. line drawing of the pectoral girdle and opercular-gular series of the holotype counterpart (ANU V2929B), c. photograph and d. line drawing of pectoral girdle, opercular-gular series and pectoral fin of the holotype (ANU V2929A). a posteroventral fragment of the operculum, a partial suboperculum, and at least seven branchiostegal rays (Fig.2). The anterior portions of most of these elements are missing, the preserved sections terminating at a margin of clean breakage, suggesting that a substantial portion of the fossil, possibly including the skull, was lost prior to collection due to weathering. The posterodorsal-most bone in the series is tentatively identified as the posterovental fragment of an operculum. It is an oblong bone bone, missing the dorsal and anterior margins. The bone surface is ornamented with short, posterolaterally directed linear ridges. The suboperculum is rectangular with a convex posterior margin. Ornament consists of short linear ridges that extend to near the posterior bone margin. At least seven branchiostegal rays are visible on ANU V2929b (Fig. 2). The first branchiostegal ray, whose dorsal margin is overlapped by the suboperculum, is more than twice as thick dorsoventrally as the other bones in the series. The 2nd ray is poorly preserved while the 3rd is narrower than the following two rays. Rays 6 and 7 are very narrow. Ornament on all bones in this series consists 40 of short rostrocaudally directed ridges with little evidence of the tubercular ornament present on the laterally facing branchiostegals of Howqualepis rostridens (Long 1988). Pectoral girdle A partial cleithrum and clavicle (Fig. 2) have a similar overall shape to those of most early actinopterygians. The cleithrum consists of an expanded ventral region with a slender vertically directed blade although the dorsal portion of this structure is missing. The bone is convex postiorly with a moderately deep embayment on the posterior margin for the insertion of the pectoral fin, similar to that of H. rostridens (Long 1988. Fig.27). The clavicle is triangular and overlaps the cleithrum posteriorly and is itself dorsally overlapped by the branchiostegal rays. Preserved sections of ornament on both the cleithrum and clavicle consists of limited areas of short ridges, particularly around the posterior margin of the clavicle and the vertical blade of the cleithrum, that are largely replaced by rostrocaudally oriented Proc. Linn. Soc. N.S.W., 130, 2009 B. CHOO rows of small pores over most of the remainder of the bone surface. This differs from the condition in Howgqualepis rostridens where the dermal surface of the corresponding area is covered ina mixture of ridges and raised tubercles with no porous ornamentation (Long, 1988. Fig.15). Donnrosenia has very similar ornamentation on the clavicle but has entirely linear ornamentation on the cleithrum (Long, Choo & Young, 2008. Fig.6). Moythomasia durgaringa and M. nitida also have porous ornamentation on the pectoral girdle, but restricted to the ventral faces of the cleithrum and clavicle (Choo, in prep) whereas pores are also present on the lateral surface in ?H. youngorum. Fins The pectoral fin (Fig. 2) is incomplete with no traces of the endoskeletal radial although the visible lepidotrichia are well preserved. The fin is elongate and triangular with more than 14 primary lepidotrichia present. As with H. rostridens and Donnrosenia, the anterior lepidotrichia are unsegmented for most of their length with secondary division restricted to the region near the fin margin. The trailing edge of the fin is not preserved and it is unclear if the posterior fin rays were fully segmented as in H. rostridens (see below). The fin reaches its maximum length at about the seventh primary ray, which is unsegmented for more than 75% of its length as in H. rostridens, longer than the c.65% unsegmented region in the fin of Donnrosenia (Long et.al, 2008). A short section of the leading edge is preserved with spine-like fringing fulcra formed by terminal branching of the leading fin rays. As with H. rostridens and Donnrosenia (see below) there is no medial contact visible between the distal hemilepidotrichia of each fringing fulcra on any of the fins. The pelvic fin (Fig. 3) is long-based and triangular. Its preserved lateral aspect and does not appear to be as elongate as in H. rostridens although it is unclear if a section of the posterior margin is missing. The fins are located approximately midway along the body between the pectoral and anal fins. Primary lepidotrichia are only preserved for the anterior half of the fin, comprising more than 22 rays suggesting more the 40 primary rays on the entire preserved section. These rays are evenly segmented along their preserved length. Slender spine-like fringing fulcra are present on the leading edge. The anal fin (Fig. 4a, b) is large and triangular in shape. At least 54 primary segmented lepidotrichia are present as opposed to c.45 fin rays on the anal fin of H. rostridens. It is unclear if the fin originally had a short posterior fringe trailing behind the main Proc. Linn. Soc. N.S.W., 130, 2009 Figure 3. ?Howqualepis youngorum sp. nov. a. pho- tograph and b. line drawing of the pelvic fin and associated squamation on ANU V2929A. triangular area of the fin as in H. rostridens. If this was the case then the complete fin would have probably had over 60 primary lepidotrichia. As in the other fins, shortened spine-like lepidotrichial segments form a serrated cutwater of fringing fulcra on the leading edge. As was the case in other known Devonian actinopterygians, the caudal fin (Fig. 4) was heterocercal in structure with a distinct posterior cleft separating the dorsal lobe (notochordal mass of the fin plus the dorsal hypochordal lobe) from the ventral hypochordal lobe. While little of its dorsal counterpart has been preserved, the ventral hypochordal lobe is elongate and triangular with c.40 primary lepidotrichia preserved. Spine-like fringing fulcra are present on the leading edge. The dorsal fin is not preserved in the holotype although a pair or large, isolated lepidotrichs preserved near the counterpart tail may have originated from that fin (Fig. 4d). Scales and squamation Articulated macromeric scales, scutes and basal fulcra are preserved from the caudal fin, extending 4] A DEVONIAN ACTINOPTERYGIAN FISH i Naeem lem Figure 4. ?Howqualepis youngorum sp. nov. a. photograph and b. line drawing of the anal and caudal fins of ANU V2929A, c. photograph and d. line drawing of the caudal fin of the holotype counterpart (ANU V2929B). forwards to above the anal fin (Fig. 4c, d). There are also isolated patches of scales preserved above and to the rear of the pelvic fins (Fig. 3). Very little of the scale ornamentation has been preserved. The visible scale types are described in accordance with the zonation terminology as proposed in in Esin (1990) and employed in Trinajastic (1999). Area C = flank scales extending from above the pelvic fins to above the anal fin. Scales are elongate and rectangular, with rostrocaudal length being at least twice the height of the scale. Ventral margin is gently convex. The disposition of the peg and socket articulation is unknown in the scales close to the pelvic fins and absent in the scales near the anal fin. Free field ornamentation is poorly preserved but individual scales show remnants of longitudinal furrows. Scales from near the front and rear of the field seem to have two or three serrations protruding along the caudal edge suggesting little or no rostrocaudal decrease in the number of serrations. Area D = scales anterior to the caudal fin and on the notochordal mass of the caudal fin. 42 Scales anterior to the caudal fin are rhombic in form, becoming smaller and increasingly elongate on the notochordal mass of the fin. Scales near area C have a gently convex ventral margin, becoming less prominent towards the caudal fin until the margin is completely straight at those scales near the caudal inversion. Peg and socket articulation is absent. The free field is smooth with no preserved traces of raised ornamentation. Posterior serrations range from two in scales near area C to none on those scales on the caudal fin. Area H = scales adjacent to the base of the anal fin. These scales are small, elongate rhomboids. Peg and socket articulation is not visible and probably absent. There is no evidence of ornamentation or posterior ridges. The only dermal scutes that have been preserved are an articulated series visible anterior to the dorsal caudal lobe and extending over the dorsal margin of the caudal fin (Fig 3b, c). Anterior to the caudal fin, the scutes are triangular plates with a caudally- Proc. Linn. Soc. N.S.W., 130, 2009 B. CHOO directed apex and are about three times longer than the adjacent flank scales. As the series progresses posteriorly over the notochordal mass of the caudal fin, the scutes narrow and spine-like with extensive overlap between the individual scutes. Redescription of the pectoral fin of Howqualepis rostridens Long (1988) described the pectoral fin of Howgqualepis rostridens as consisting of 25 primary lepidotrichia that are unsegmented for most of their extent, save for some secondary division near the fin margin. A complete pectoral fin was not figured and re-examination of this form has revealed the fin to be more extensive than previously recognised (Fig. 5). Additionally, the leading edge of the pectoral and other fins was described as having short, parallel rays similar to fringing fulcra, but not paired (ibid). A similar condition in Donnrosenia led to Long et. al (2008) to diagnose the Howqualepididae as possessing short spine-like lepidotrichia in lieu of true fringing fulcra. The anterior two-thirds of the fin consist of c.25 lepidotrichia that possess extensive proximally unsegmented sections that in some specimens display distal bifurcation. At the lateral margins, these primary rays branch into a fringe of narrow, segments. The relative length of the proximal rays to the segmented 5mm f.ful fringe is variable, with the unsegmented region accounting for between 75-90% of the length of the fin. There appears to be no correlation between the degree of distal segmentation and the size of the specimen. Posterior of the unsegmented rays are at more than 10 additional primary lepidotrichia that are segmented from base to margin, again displaying a variable degree of distal branching. The pectoral fin of H. rostridens was thus broader in shape and less-extensively unsegmented than has previously been described. In the majority of specimens, the delicate elements of the posterior rays and terminal fringe are scattered or missing, leaving only the thick unsegmented proximal sections in articulation. This configuration of the pectoral fin-rays is similar to that of a number of Carboniferous taxa including Rhadinichthys (Moy-Thomas & Bradley Dyne, 1938). On the leading edge of the pectoral fins of Howgqualepis rostridens, ?H. youngorum sp.nov and Donnrosenia, the terminal sections of the otherwise unsegmented marginal fin rays branch at least twice, to forming narrow spine-like elements that are not obviously paired. These elements are called “terminal lepidotrichia” in Cheirolepis (Pearson and Westoll, 1979) and Melanecta (Coates, 1998) or “cutwater lepidotrichia” in the Howqualepididae Figure 5. Pectoral fin of Howqualepis rostridens. a. photograph and b. line drawing of the fin of MV P.160857. c. photograph and d. line drawing of the fin of MV P.160851B. In this specimen, the posterior section has partially torn off and folded to be visible ventral of the anterior edge of the fin. Proc. Linn. Soc. N.S.W., 130, 2009 43 A DEVONIAN ACTINOPTERYGIAN FISH (Long, Choo and Young, 2008). In a recent study, such structures fall into Arratia’s “Pattern A” class of fringing fulcra, formed from overlapping branched projections of the anteriormost lepidotrichia (Arratia, in press), a condition found in all undisputed Devonian actinopterygians with the exception of Tegeolepis which appears to lack any sort of spiny cutwater (Dunkle and Schaeffer, 1973). The fulcra of Cheirolepis, which are of similar form to those of the Howqualepididae, comprise distally enlarged hemilepidotrichia that partially enclose their paired counterparts (Arratia, in press). The more obviously paired structures present in Moythomasia and “Mimia” (also falling within “Pattern A”) are the result of the terminal segments being of equal length and in medial contact. Given that the scheme proposed by Arratia (and adopted here) means that all Devonian fringing fulcra are in fact modified spine- like lepidotrichia (merely differing in the nature of contact between the hemilepidotrichia), the diagnosis of Howqualepidiae has been adjusted accordingly in the systematic description. DISCUSSION Long, Choo and Young (2008) erected the Howqualepididae, comprising Howqualepis rostridens from Mount Howitt, Victoria and Donnrosenia schaefferi from the Aztec Siltstone of Antarctica. ANU V2929 appears to represent a third taxon within this clade (Fig. 6). All three fish have an elongate body form with macromeric squamation; long-based pelvic fins; small fringing fulcra without medial contact between the distal hemilepidotrichia, and extensive unsegmented primary lepidotrichia that comprise most of the length of the pectoral fin. Among the other Devonian actinopterygians, only 7egeolepis clarki (Dunkle and Schaeffer, 1973) possesses extensive unsegmented pectoral lepidotrichia but is distinguished from the Gondwanan forms in lacking a terminal segmented fringe on the pectoral fins, in possessing micromeric squamation and having small, short-based pelvic fins. Assigning the Bunga Bed taxon to a genus is rendered difficult owing to the lack of key skull characters that are used to characterise Howqualepis rostridens from the similar Donnrosenia. For example, H. rostridens possesses an extremely long maxillary blade, a dentigerous rostral and small, dorsoventrally compressed premaxillae (Long 1988). Donnrosenia displays a short, deep maxillary blade, dorsoventrally prominent premaxillae, a small accessory operculum and much smaller teeth than Howqualepis (Long, 44 Choo and Young, 2008). ANU V2929 is considered to be closer H.rostridens in having more extensive unsegmented pectoral lepidotrichia and relatively smaller scales than Donnrosenia. The pectoral fins of ANU V2929 are more similar to that of H. rostridens in that both forms possess unsegmented lepidotrichia that account for over 75% of the maximum length of the fin. Those of Donnrosenia account for less than 70% of the maximum fin length (Long, Choo and Young, 2008. Hige7): Based on these anatomical similarities and pending the discovery of skull material for this taxon, ANU V2929 is tentatively assigned to Howqualepis. The Bunga Bed form is not conspecific with H. rostridens and is distinguished in having a larger anal fin with a greater number of primary lepidotrichia and in possessing porous dermal ornamentation of the pectoral girdle. The presence of a grade of Devonian actinopterygian so far found exclusively in Middle Devonian freshwater deposits of southeastern Australia and Victoria Land, Antarctica highlights the close biogeographical similarity between the fossil faunas of these two regions. The apparent absence of these ray-finned fishes in Devonian sites outside this area also adds to a growing body of fossil evidence that indicates a regionally endemic freshwater vertebrate fauna within Middle Devonian Eastern Gondwana. Similarities in key taxa of placoderms (Young 1988, Young and Long 2005), acanthodians (Long 1983, Young 1989, Young & Burrow 2004), chondrichthyans (Young 1982, 2007; Long & Young 1995) and dipnoans (Long 1992, 2003) have been well documented. ACKNOWLEDGEMENTS Thanks to Gavin Young and John Long for their supervision and helpful suggestions regarding this manuscript. Thanks also to Gloria Arratia for helpful discussion regarding fringing fulcra. Ben Young is commended for his excellent fossil preparation and photography. Proc. Linn. Soc. N.S.W., 130, 2009 B. CHOO Se os SS SSS SSS TN AS SS C4 Ceri pemyh— tt Sa 7] ASO Sy SSeS SESE SS a SSS SSS SS SSSA SEES cate Wat tere TAA Seewecween= Lis — mest: Sas tt cr ek wee Seo REL SoS LL a ch EEEREnawenensns= sy Ake Se SENS mm Figure 6. Comparison of the three known species of the Howqualepididae. Reconstructions presented in lateral view and are not to scale. Unknown parts of the anatomy are represented by dark grey areas. a. ?Howqualepis youngorum sp.novy., based on the preserved extent of the holotype with outline based on H. rostridens, c.14cm long. b. Howqualepis rostridens from Mount Howitt, Victoria (modified after Long, 1988). Size of specimens range from 20-50cm. c. Donnrosenia schaefferi from the Aztec Siltstone, South- ern Victoria Land, Antarctica (from Long et.al, 2008), c. 14cm long. REFERENCES Arratia, G. (in press). Identifying patterns of diversity of the actinopterygian fulcra Acta Zoologica (Stockholm). Basden, A.and Young, G.C. (2001). A primitive actinopterygian neurocranium from the Early Devonian Taemas Formation, Burrinjuck area, New South Wales, Australia. Journal of Vertebrate Paleontology, 21, 754-766. Proc. Linn. Soc. N.S.W., 130, 2009 Basden, A., Young, G.C., Coates, Mand Ritchie,A. (2000). The most primitive osteichthyan braincase? Nature, 403, 186-188. Burrow, C. J. (1996). Taphonomy of acanthodians from the Devonian Bunga Beds (late Givetian — early Frasnian) of New South Wales. Historical Biology 11, 213-228. Cas R. A. F., Edgar C., Allen R. L., Bull S., Clifford B. A., Giordano G. and Wright J. V. (2000). Influence of magmatism and tectonics on sedimentation in 45 A DEVONIAN ACTINOPTERYGIAN FISH an extensional lake basin: the Upper Devonian Bunga Beds, Boyd Volcanic Complex, south- eastern Australia. International Association of Sedimentologists Special Publications 30, 175-200. Choo, B., Long, J.and Trinajstic, K. (in press). A new genus and species of basal actinopterygian fish from the Upper Devonian Gogo Formation of Western Australia. Acta Zoologica (Stockholm). Coates, M. I (1998). Actinopterygians from the Namurian of Bearsden, Scotland, with comments on early actinopterygian neurocrania. Zoological Journal of the Linnean Society 122, 27-59. Cope, E. D. 1887. Geology and Palaeontology. American Naturalist, 1014-1019. Dunkle, D.and Schaeffer, B. (1973). Tegeolepis clarki (Newberry), a palaeonisciform from the Upper Devonian Ohio Shale, Palaeontographica Abteilung A 143, 141-158. Esin, D.N. (1990). The scale cover of Amblypterus costata (Eichwald) and the palaeoniscid taxonomy based on isolated scales. Paleontological Journal 2,90-98. Fergusson C. L., Cas R. A. F., Collins W. J., Craig G. Y., Crook K. A. W., Powell C. McA., Scott P. A. and Young G. C. (1979). The Late Devonian Boyd Volcanic Complex, Eden, N.S.W. Journal of the Geological Society of Australia 26, 97-105. Gardiner, B.G. (1984). The relationships of the palaeoniscoid fishes, a review based on new specimens of Mimia and Moythomasia from the Upper Devonian of Western Australia. Bulletin of the British Museum (Natural History) Geology, 37, 1-428. Gardiner, B.G. and Bartram, A.W.H. (1977). The homologies of ventral cranial fissures in osteichthyans, pp. 227-245. In S. M. Andrews, R. S. Miles and A. D. Walker (eds.), Problems in Vertebrate Evolution. Academic Press, London. Giordano G.and Cas R. A. F. (2001). Structure of the Upper Devonian Boyd Volcanic Complex, south coast New South Wales: implications for the Devonian — Carboniferous evolution of the Lachlan Fold Belt. Australian Journal of Earth Sciences 48, 49-61. Huxley, T. H. (1880). On the applications of the laws of evolution to the arrangement of the Vertebrata and more particularly of the Mammalia. Proceedings of the Zoological Society of London, 1880, 649-662. Jessen, H. (1968). Moythomasia nitida Gross und M. cf. striata Gross, Devonische palaeonisciden aus dem oberen Plattenkalk der Bergish-Gladbach- Paffrather Mulde (Rheinisches Schiefergebirge). Palaeontographica Abteilung A 128, 87-114. Long, J.A. (1988). New palaeoniscoid fishes from the Late Devonian and Early Carboniferous of Victoria. Memoirs of the Association of Australian Palaeontologists, 7, 1-64. Long, J. A.; Choo, B.and Young, G. (2008).A new basal actinopterygian fish from the Middle Devonian Aztec Siltstone of Antarctica. Antarctic Science 20, 393- 412. 46 Moy-Thomas, J. A.and Bradley Dyne, M. (1938) The actinopterygian fishes from the Lower Carboniferous of Glencartholm, Eskdale, Dumfriesshire. Transactions of the Royal Society of Edinburgh, 59, 437-480. Pearson, D. M.and Westoll, T. S. (1979) The Devonian actinopterygian Cheirolepis Agassiz. Transactions of the Royal Society of Edinburgh 70, 337-399. Rickard M. J. and Love S. (2000). Timing of megakinks and related structures: constraints from the Devonian Bunga-Wapengo Basin, Mimosa Rocks National Park, New South Wales. Australian Journal of Earth Sciences 47, 1009-1013. Schultze, H.-P. (1968). Palaeoniscoidea-Schuppen aus dem Unterdevon Australiens und Kanadas und aus dem Mitteldevon Spitzbergens. Bulletin of the British Museum (Natural History): Geology, 16, 342-368. Trinajstic, K. (1999). Scale morphology of the Late Devonian palaeoniscoid Moythomasia durgaringa Gardiner and Bartram, 1997. Alcheringa, 23, 9-19. Young, G.C. (1982). Devonian sharks from south-eastern Australia and Antarctica. Palaeontology 25, 817-843. Young, G.C. (1999). Preliminary report on the biostratigraphy of new placoderm discoveries in the Hervey Group (Upper Devonian) of central New South Wales. Records of the Western Australian Museum, Supplement 57, 139-150. Young, G. C. (2007). Devonian formations, vertebrate faunas and age control on the far south coast of New South Wales and adjacent Victoria. Australian Journal of Earth Sciences 54, 991-1008. Proc. Linn. Soc. N.S.W., 130, 2009 Fire and Habitat Interactions in Regeneration, Persistence and Maturation of Obligate-seeding and Resprouting Plant Species in Coastal Heath PETER J. MYERSCOUGH Institute of Wildlife Research, School of Biological Sciences, The University of Sydney, Sydney, NSW 2006. Email: pmyersco@bio.usyd.edu.au Myerscough, P.J. (2009). Fire and habitat interactions in regeneration, persistence and maturation of obligate-seeding and resprouting plant species in coastal heath. Proceedings of the Linnean Society of New South Wales 130, 47-61. After a fire in January 1991, populations of two obligate-seeding and two resprouting species were followed from seeds sown in dry heath and wet heath on Pleistocene beach sands in the Myall Lakes area. In each type of heath, there were four plots, each with ninety 25 X 25 cm quadrats in which seeds of the four species had been sown in various combinations and surface soil conditions. All four wet-heath plots burned again in January 1998, as did two of the dry-heath plots. The two obligate-seeding species were confined to their respective habitats early in the life cycle; Acacia ulicifolia to dry heath by lack of seeds and suitable conditions for seedling emergence in wet heath; Dillwynia floribunda to wet heath by failure of its seedlings to survive in dry heath. The two resprouting species were confined to their respective habitats in different ways; Banksia oblongifolia by failure of its seedlings to survive in dry heath; Banksia aemula by lack of suitable soil surface in wet heath for establishment of its seedlings. In both species of Banksia, seedlings require a lignotuber to survive their first fire, and may persist several years without appreciable growth. Manuscript received 11 August 2008, accepted for publication 17 December 2008 KEY WORDS: banksias, fire, heath, lignotubers, maturation, oskars, persistence, regeneration, resprouters, seeders INTRODUCTION Dispersal, survivalandreproductionofindividuals underlie patterns of distribution and abundance of species. Fire influences these processes in plant life histories, and moulds patterns evident in fire-prone vegetation across gradients in habitat. In fire-prone vegetation, species of seed plants tend to fall into two groups (Gill 1981), obligate-seeders, those whose adult plants die in fires that destroy their leaf canopies and regenerate after fire solely from seed, and resprouters, some of whose plants survive complete loss of their canopy in intense fires and resprout new canopies after fire from vegetative tissue that is protected from fire. Frequent fires may act selectively and reinforce the respective characteristics of these two groups of plants. In obligate-seeders, high production of seeds, in amount and early availability after fire, would be expected, with the seeds protected from burning, either by being contained in fire-resistant fruits or by dispersal to safe sites in soil and having dormancy that is only readily broken by stimuli connected with the passage of fire. In resprouters, seedlings would be expected to produce at an early stage vegetative parts that survive fire. Pate et al. (1990) showed that seedlings of obligate-seeding species devote much growth to their shoots and early seed production, while seedlings of comparable resprouting species devote a high proportion of their growth to underground tissues including fire-resistant vegetative storage organs. In seedlings of resprouters, production of fire-resistant vegetative tissue typically precedes seed production. In their life histories, seed production is usually considerably delayed compared with related obligate-seeding species. Obligate seeders probably have simpler relationships linking seed dispersal, germination and seedling establishment in particular environments, their regeneration niches (sensu Grubb 1977), to seed production than do resprouters. Resprouters, while passing through seed dispersal, germination and seedling establishment in particular environments, their regeneration niches, also have periods of persistence as vegetative plants, that though fire-hardy, FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS may or may not become reproductive and produce seed. It is possible that resprouters may simply persist many years as small plants with little net growth. The persistence niche (sensu Bond and Midgley 2001) of resprouters may be wider than conditions under which the plants progress to seed production. The opportunity arose to observe through time seedlings of obligate-seeding and resprouting species after fire occurred across habitats in fire-prone coastal heath. Such heath occurs in south-eastern Australia on leached siliceous beach sands and dunes deposited during the Pleistocene from South Australia (Specht 1981) to sand islands such as North Stradbroke Island (Clifford and Specht 1979) off the south-eastern coast of Queensland. On the coast of New South Wales, they occur particularly north of Newcastle to the Queensland border (Griffith et al. 2003, Keith 2004). In the Myall Lakes area, heath occurs on a Pleistocene system of beach sands in the Eurunderee Embayment of Thom et al. (1992). On these sands, there is a catenary sequence of soils and vegetation with dry heath on ridges, wet heath on slopes and swamps in periodically waterlogged swales (Carolin 1970, Myerscough and Carolin 1986, Myerscough et al. 1995). Dry heath belongs to the Banksia _ serratifolia (aemula) Alliance of Beadle (1981) and Wallum Sand Heaths of Keith (2004), and wet heath to Beadle’s (1981) Banksia aspleniifolia (oblongifolia) Alliance and Keith’s (2004) Coastal Heath Swamps. Fire has occurred fairly frequently, but over two decades produced no detectable effect in changing the pattern of differentiation of vegetation across the sequence of habitats, though changes with time since fire were clearly evident in the vegetation within habitats (Myerscough and Clarke 2007). Carolin (1970) demonstrated that various species occupy characteristic ranges of habitat in the catenary sequence from the ridges to the swales. Myerscough et al. (1996) and Clarke et al. (1996) investigated in four species how occupancy of their ranges of habitat might arise through dispersal of seed and, after fire, germination and establishment of their seedlings. Seedlings of two species characteristic of wet heath, obligate-seeding Dillwynia floribunda and resprouting Banksia oblongifolia, did not survive in dry heath, despite their seeds occurring and germinating there (Myerscough et al. 1996). Seed of two species characteristic of dry heath, obligate-seeding Acacia ulicifolia and resprouting Banksia aemula, were at best rare in wet heath (Myerscough et al. 1996), and, unless the soil surface is artificially disturbed and seeds are buried, germination and seedling establishment did not occur (Clarke et al. 1996). In short, it was largely in regeneration niche (Grubb 1977) that these species 48 appeared to be segregated to their respective habitats, D. floribunda and B. oblongifolia to wet heath, and A. ulicifolia and B. aemula to dry heath (Myerscough et al. 1996, Clarke et al. 1996). Seedlings of the four species were observed beyond the phase of establishment. Establishment of seedlings of Acacia ulicifolia and Banksia aemula had occurred in wet heath, following experimental manipulation of the soil surface (Clarke et al. 1996). Early survival of seedlings of both obligate-seeding species was related to type of habitat, but in both resprouting species it was related to variation among plots within types of habitat (Clarke et al. 1996). In this paper, ongoing survival of seedlings of the two obligate-seeding species is examined in relation to type of habitat, while in the two resprouting species it is examined in relation to variation among plots within types of habitat. Fire recurred in six of the eight experimental plots seven years after the fire that immediately preceded the start of the experiment. Survival of seedlings of the two resprouting species through fire could thus be assessed. Benwell (1998) observed lignotubers in seedlings of Banksia aemula and B. oblongifolia in similar coastal heath and found their growth to be slow seven years after fire. Four years after our experiment started, lignotubers were observed on some seedlings of each of the two species. By using fire-proof tags and measuring sizes and positions of lignotubers, survival of seedlings through their first fire could be assessed in relation to size and position of their lignotubers, if indeed they had been formed. Auld (1987) had found in Angophora hispida that seedlings with buried lignotubers survived fire better than those lacking lignotubers or with them exposed above the soil surface. Ongoing observation of lignotubers and sizes of banksia seedlings was used to try to identify whether they were growing or merely surviving without net growth. One seedling of Banksia aemula was observed to flower and set fruit. It was thus possible to see whether a fire-proof stem was required for flowering and seed production as Bradstock and Myerscough (1988) found in juveniles of Banksia serrata. The questions investigated in this paper are: * Do patterns of early seedling survival in the two obligate-seeding species seen in relation to type of habitat continue into later stages, and how are these patterns related to flowering and seeding? * How are patterns of seedling survival in the two resprouting species related to characteristics of individual plots? ¢ What roles do formation, size and position of lignotubers play in the survival through fire of Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH Table 1. Experimental plot locations and their transect, ridge position relative to coastline, habitat, and fire history since January 1991. Habitat Fire history Jan 1998 Nov 2006 WH Totally burnt Totally burnt WH Totally burnt Totally burnt Totally burnt but WH some scorched Totally burnt leaves present Totally burnt but Burnt but with WH some scorched some patches leaves present unburnt DH Totally burnt Totally burnt DH Totally burnt Totally burnt DH Unburnt Unburnt Plots GDA Transect Ridge 82° S29 E 2975S. T2W1 1125 2 Near ASH. S T2W3 21013 E T2 Far DO MOZS.° T3W1 22310 E T3 Near 29.083 S* T3W2 12.250'E T3 Mid 29.594 S* T2D2 71.040'E T2 Mid 29.500'S T2D3 71.026 IZ, Far DSS 4 T3D1 12302 E T3 Near 29.019'S ’ 32 22. 280F T3 Mid Most or less DH unburnt - one edge slightly scorched Mostly unburnt — some lightly scorched patches * 30 X 5-m plot extends to left of marker post when facing inland; other plots extend to right of post. Dry heath (DH) and wet heath (WH). seedlings of the two resprouting species? * Do seedlings of the two resprouting species show appreciable net growth, and under what conditions may they do so? * Do patterns of seedling growth and survival give evidence of the longterm stability of the patterns observed in the vegetation across habitats of this coastal heath? METHODS AND MATERIALS Study area The heath studied was on sands of a Pleistocene beach system in the Euruderee Embayment of Thom et al. (1992). Twenty-four plots, 3 wet-heath and 3 dry-heath plots in each of 4 transects, were used by Myerscough et al. (1995) to analyse floristic variation in heaths across the system. Each plot was 30 X 5m with its longer sides parallel to the nearest beach ridge. Eight of the plots, two wet-heath and two dry-heath plots on each of the two central transects, were used Proc. Linn. Soc. N.S.W., 130, 2009 in the experiments of Myerscough et al. (1996) and Clarke et al. (1996). Each of these plots (Table 1) was divided into a grid of 150 square-metre cells. Ninety cells were randomly chosen and to each of these cells a 25 X 25 cm quadrat was randomly allocated to a particular experimental treatment. Experimental treatments, including placement of seeds of the four species of this study, are described in Myerscough et al. (1996). These ninety quadrats in each of the 8 plots were the areas in which seedlings that arose in 1991 were observed. Data collection Periodic counts of seedlings of Acacia ulicifolia and Dillwynia floribunda were maintained from 1991 until the fire of January 1998 burned six of the eight plots (Table 1). Between 1995 and 1997, due to the density of stems, especially in wet heath, it became increasingly difficult to count seedlings of D. floribunda and A. ulicifolia on the small quadrats. Since there was no seedling recruitment apparent during this period, where a greater number of seedlings 49 FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS on a plot was recorded six months after the previous count, the greater number was taken to be correct. After January 1998 until October 2008, individuals of Acacia ulicifolia continued to be counted on the unburnt plots T3D1 and T3D2. In Banksia aemula and B. oblongifolia, all survivors of the 1991 cohort of seedlings were counted on the eight plots. In November 1995, surviving banksia seedlings were marked with fireproof metallic tags on stainless steel pins placed beside the seedlings. All seedlings of Banksia aemula were tagged. In B. oblongifolia, many more seedlings were then surviving, and in those quadrats where there was more than one seedling only one seedling in the 25 X 25 cm quadrat was randomly selected and tagged. The proportion of individuals tagged in November 1995 and the number of tagged individuals subsequently surviving were used to estimate the population of surviving seedlings of B. oblongifolia in each plot. When no tagged individuals had survived in a plot, it was assumed that the whole cohort of seedlings that had arisen in 1991 in the experimental quadrats of the plot had died. Lignotuber development was followed on each of the tagged seedlings, noting whether a lignotuber was absent or present. If present, its mean width was recorded from two measurements taken in two directions at right angles, and if it was not entirely buried, the height of its top above the soil surface was measured. After the fire of 1 January 1998, in March 1998 survival of the tagged seedlings was assessed. A seedling was scored as dead if it failed to resprout and live if it had resprouted. Most seedlings that resprouted had done so by March 1998, but a few resprouted later and were identified as alive when scored some months later. In all tagged seedlings, alive or dead, lignotuber presence or absence was noted, and, if present, its mean width was measured and whether its top was buried or exposed. The top was scored as exposed if its height above the soil surface was greater than | mm. Survival of seedlings through the fire was assessed in relation to habitat and lignotuber presence and exposure above the soil using 2 X 2 contingency tables and Chi square statistic. Growth of banksia seedlings between 1995 and 2007 was assessed from lignotuber width and plant height. In Banksia aemula, seedlings were deemed to have grown if in October 2007 they were found to have a lignotuber width of over 40 mm or a plant height of greater than 40 cm, while in Banksia oblongifolia seedlings with a lignotuber width of over 20 mm were deemed to have grown. Widths of lignotubers of Banksia aemula were not easily assessed in a consistent way through time for two reasons. Firstly, 50 although two measurements of width taken at right angles to each other were made on each occasion, not all lignotubers are radially symmetrical. Secondly, the lignotubers form with a thick bark, as in the sister species Banksia serrata (Beadle 1940, Bradstock and Myerscough 1988), and this bark may erode so that measured widths of lignotubers may lessen in time. Thus it is possible that some of the seedlings of Banksia aemula deemed not to have grown between 1995, or from when their lignotuber formed if it was later than 1995, may actually have grown slightly. Watertables were observed in the plots between 1991 and 1997, and their depths recorded as described in Myerscough et al. (1996). The fire of January 1998 prevented further observations, destroying tops of the plastic pipes used to observe depths to the watertable on 6 of the 8 plots. The depths given in Table 2 were measured on 23 September 1997, when the watertable was relatively high. To illustrate key floristic variation observed in 1990 across the plots, the nineteen most abundant species were selected from Appendix II of Myerscough et al. (1995) and listed in Table 2 in the order in which they were sorted in the TWINSPAN analysis given in Appendix I of Myerscough et al. (1995). The nineteen species included Banksia aemula, B. oblongifolia and Dillwynia floribunda. The other species, Acacia ulicifolia, whose seedlings were observed on the experimental plots was also included. The height of the canopy of each of the plots was recorded in September 2005 in ten randomly selected 1 X 1 m cells, except in T2W3 where inadvertently there were only nine cells. In each cell, the species of the tallest plant was noted. At the same time, the degree to which each surviving banksia seedling was shaded by surrounding vegetation was subjectively scored using a five-point scale of shade: 5, >95%; 4, 95-75%; 3, <75-25%; 2, <25% shaded; 1, seedling’s canopy unshaded. Nomenclature Nomenclature of plant names used follows Harden (1990, 1992, 1993 and 2002). RESULTS The plots differed floristically and in depths to the watertable (Table 2). Depths to watertable were greater in dry heath than in wet heath plots (F, =5.90 (p just >0.05)) and differed markedly among plots within habitats (F,,,=299.3 (p<0.001)). The habitats differed in plant species that provide significant cover. Both habitats had shrubs with appreciable Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH Table 2. Experimental plots and mean (SE) depth (cm) to watertable and mean cover (“%) of twenty species (RS, resprouter; OS, obligate-seeder). Plot T2W1 T2W3 T3W1 T3W2 T2D2 T2D3 SIDI T3D2 2.0 10.6 2.8 18.9 43.3 26.3 108.8 38.5 Watertable OD CI CA" MIDE NOS MOGY OMOAR OS) Empodisma minus RS Gymnoshoenus sphaerocephalus 29 RS Leptospermum livesidgei RS 44 Banksia oblongifolia RS Dillwynia floribunda OS Epacris obtusifolia OS Xanthorrhoea fulva RS Lepyrodia interrupta RS 11 15 34 18 3 4 59 36 16 Darwinia leptantha OS Pseudanthus orientalis RS Persoonia lanceolata OS Kunzea capitataOS 1 9 5 14 ; 1 Dillwynia retorta OS Leptospermum polygalifolium RS 49 25 3 Leptospermum trinervium RS Acacia ulicifolia OS Banksia aemula RS 1 20 52 26 23 Melaleuca nodosa B RS Hypolaena 10 Vi 1S 4 fastigiata RS Epacris pulchella OS Proc. Linn. Soc. N.S.W., 130, 2009 5] FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS cover such as the banksias, Banksia aemula in dry heath and B. oblongifolia in wet heath. Wet heath had more cover from resprouting monocotyledons such as Xanthorrhoea fulva than dry heath, and more cover from obligate-seeding shrubs such as Dil/lwynia floribunda and Epacris obtusifolia with sparsely branched, elongate ascending stems. Though similar in depth to the watertable, the wet heath plots T2W1 and T3W1 differed in plant cover. T2W1 had high cover of Empodisma minus and Gymnoschoenus sphaerocephalus. After the fire of January 1991, and sowing seeds in March 1991 under various treatments across the eight plots, as described in Myerscough et al. (1996), seedlings of Banksia aemula, B. oblongifolia, Acacia ulicifolia and Dillwynia floribunda differed in their patterns of survival across the plots (Table 3). In dry heath plots, seedlings of Dillwynia floribunda, though fairly numerous at six months, suffered heavy mortality and were completely absent after four years. They persisted in all wet heath plots with approximately 10% of the population observed at six months present six years later, with some plants observed to have flowered after three and half years. All the plants in the plots were killed by the fire of January 1998. In short, it was only in the wet heath plots that plants of D. floribunda survived and reproduced, doing so with little plot to plot variation apparent in their survival (Table 3). Seven and a half years after the fire in January 1998, D. floribunda was among emergent species in the canopy of the wet heath plots (Table 4). Some seedlings of Acacia ulicifolia survived from 1991 in each of the eight plots until the fire of January Table 4. Experimental plots and height (m) of canopy (mean (SE)), emergent species and relative shading (*RSh) of banksia seedlings (numbers in each category) in September 2005. Plot T2W1 T2W3 T3W1 T3W2 T2D2 T2D3 T3D1 T3D2 Ganon erent 1.50 e337) 1.38 1.09 2.08 1.93 DMS 1.63 (0.06) (0.06) (0.06) (0.07) (0.14) (0.16) (0.16) (0.08) @ Emergent Dif 3 Dfl2 Daria Dfl2 B.ae 3 B.ae4_ Bae4 B.ael Species - number TiS Io.15 1 JET JEG Ml te S LL GAM L.tr 4 L.tr 4 of contacts out of S.in 3 S.in 2 S.sp 1 D.re 2 10 (but out of 9 for Piaal las Pla3 T2W3) A.el | L.po | E.mi | E.mi 1 E.ob 2 E.mil B.ob 1 B.ob 1 B.ob 1 A.te | B fal Crem Mn | Wp | X fu | K.ca | B. aemula RSh 5 3 4 4 3 9 5 2 2 3 3 4 5 Sil 2 5 2 1 2 10 1 1 5 1 1 1 B. oblong- Rsh 5 folia 4 2 2 3) 1 1 1 2 2 1 @ A.el — Acacia elongata; A.te — Acacia terminalis; B.ae — Banksia aemula; B. ob — Banksia oblongifolia; B.fa— Boronia falcifolia; C.te — Calytrix tetragona; D.fl— Dillwynia floribunda; D.re — Dillwynia retorta; E.mi — Epacris microphylla; E.ob — Epacris obltusifolia: K.ca - Kunzea capitata; L.li— Leptospermum liversidgei; L.po — Leptospermum polygalifolia; L.tr — Leptospermum trinervium: M.n — Melaleuca nodosa; P.la— Persoonia lanceolata; S.in — Sprengelia incarnata; S.sp — Sprengelia sprengelioides; W.p — Woollsia pungens; X.fu — Xanthorrhoea fulva. * RSh: 5, >95%; 4, 95-75%; 3, <75-25%; 2, <25% shaded; 1, seedling’s canopy unshaded. 52 Proc. 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Linn. Soc. N.S.W., 130, 2009 FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS 1998 burned six of the plots. Though survival varies considerably with plot, there is no clear pattern in this variation in relation to habitat or other characteristics of plots. In the two plots not burned in 1998, one plant continued to survive in T3D2 until it was fifteen and a half years-old, while in T3D1 five plants were still alive at 17.65 years (Table 3), four of them having fruited in 2008. In this plot, four-year-old plants flowered and fruited, and four-year-old plants were seen flowering on other plots (T2D3 and T2W3). In Banksia oblongifolia, some seedlings survived on each of the eight plots up to two years (Table 3). On dry heath plots, they had died out after 5 years on both plots where the watertable was deep (T2D2 and T3D1) but continued to survive in significant number on T2D3, the dry heath plot with the least depth to the watertable (Table 2). No seedling of B. oblongifolia survived the fire of January 1998 on a dry heath plot, but on each of the four wet heath plots some seedlings survived. On T2W1, the plot with high cover of Gymnoschoenus sphaerocephala and Empodisma minus (Table 2), no seedling survived twelve years, but on the other three wet heath plots some seedlings survived up to seventeen years (Table 3). In Banksia aemula, some seedlings survived on each of the eight plots up to nine and half years, including on the six plots totally burnt in the fire of January 1998. Their numbers were lowest on the two dry heath plots (T2D2 and T3D1) where the watertable was deep and cover of Leptospermum trinervium relatively high (Table 2). On T3D1, which had the deepest watertable (Table 2) and which was not burnt in 1998 (Table 1), the last survivor had died after ten years. On each of the other seven plots, at least one plant survived to seventeen years (Table 3). Among wet heath plots, there was heavier mortality of survivors of the fire of January 1998 on T2W1 and T3W1 (see years 7.64 to 17.65 in Table 3), plots with the shallowest watertable and highest cover of resprouting monocots (Table 2), than on T2W3 and T3W2. On T2W3 and T3W2, not only was the watertable deeper and the cover of monocots less (Table 2), but in September 2005 the surviving seedlings of Banksia aemula were less shaded (Table 4). There was one seedling of B. aemula on each of these plots that was unshaded (Table 4). On T2W3, one plant flowered at fourteen years and formed swollen follicles, and, after the fire in November 2006, six follicles appeared to have opened. This was the only banksia originating from seed in 1991 that was observed on any of the eight plots to have become reproductive. Across the wet heath plots, mortality from the fire of | January 1998 was much higher among seedlings of Banksia oblongifolia (83%) than among those of B. aemula (23%) (p<0.001). In both species of banksia, survival of seedlings on plots burnt in the fire of 1 January 1998 entirely depended on possessing a lignotuber; without a lignotuber no seedling survived (Table 5). Under comparable conditions in wet heath, the lignotubers of B. aemula survived better than those of B. oblongifolia. With the top of the lignotuber exposed, only 10% of seedlings of B. oblongifolia survived whereas 78% of those of B. aemula survived; with the lignotuber buried, 36% survived in B. oblongifolia and 95% in B. aemula. No tagged seedling of B. oblongifolia survived fire in a dry heath plot (Tables 3 and 5), while seedlings of B. aemula survived fire in both wet heath (WH) and dry heath (DH). The survival of B. aemula seedlings was much lower in DH (24%) than in WH (76%) not only because there was a higher proportion of seedlings without lignotubers in DH (24%) than in WH (7%) (p<0.001) but there was higher mortality of seedlings with lignotubers in DH (68%) than in WH (18%) (p<0.001). Burial of the lignotuber Table 5. Number of tagged banksia seedlings live or dead in March 1998 after fire of 1 January 1998 in relation to habitat and lignotubers. Species Banksia aemula Banksia oblongifolia Habitat Dry heath Wet heath Dry heath Wet heath Seedlings Live Dead Live Dead Live Dead Live Dead Lignotuber: absent 0 14 0 8 0 3) 0 5 present 14 30 93 21 0 10 19 117 Lignotuber top: buried 11 21 20 I 0 2 8 14 exposed 3) 9 We 20 0 8 1] 103 54 Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH Table 6. Dimensions and relative shading (RSh) of grown tagged banksia seedlings in October 2007. Species Banksia aemula Banksia oblongifolia Dimension Mean Plant Relative Mean Plant Relative lignotuber height shading lignotuber height shading width (mm) (cm) (RSh)@ ~~ width (mm) (cm) (RShH)@ Dry heath plot T3D2 4] 43 3 Wet heath plots Al 14 2 3 16 2 T2W3 43 Sil 2 39 28 2) 74* Ue ] 33 76 ] T3W2 37) 72 1 * Plant first flowered in 2005. @ RSh: 1, seedling’s canopy unshaded; 2, <25%; 3, <75-25% shaded increased the chances of survival of seedlings, particularly in B. oblongifolia. In B. aemula, the extent of this was mediated by habitat. A higher proportion of lignotubers were buried in DH (73%) than in WH (18%) (p<0.001). Despite this, mortality of seedlings with buried lignotubers was much higher in DH (66%) than in WH (5%) (p<0.001), whereas seedlings with lignotubers exposed above ground suffered 75% mortality in DH and 22% mortality in WH (p<0.001). In short, though burial of their lignotubers enhanced survival of seedlings in both habitats, it was more effective in WH than DH though the proportion of seedlings with buried lignotubers was lower in WH than DH. Of those tagged banksia seedlings surviving to October 2007, appreciable growth was detected in relatively few (Tables 6 and 7), and most of these seedlings occurred in one wet heath plot, T2W3. Indeed, in this plot, two of the three surviving seedlings of Banksia oblongifolia, and four of the six surviving seedlings of Banksia aemula had grown, with one of them flowering in 2005 and producing an infructescence with a single swollen follicle. This individual was the only seedling to have had a lignotuber over 40 mm in width by March 1998; no others had achieved this by September 2005. In March 2007, it had four infructescences on which a total of six follicles had opened after the fire in November 2006. This was the only tagged banksia Proc. Linn. Soc. N.S.W., 130, 2009 seedling to have reached reproductive maturity. In all the other plots, there were only two tagged banksia seedlings that could be identified as having grown, both B. aemula, one on a dry heath plot, T3D2, and the other on a wet heath plot, T3W2. The rest of the surviving tagged banksia seedlings appeared to be simply surviving without net growth, and on the wet heath plot T3W2 such seedlings of B. aemula were particularly numerous (Table 7). In October 2007, all seedlings deemed to have grown were unshaded or <25% shaded (Table 6), except for the seedling on T3D2, a plot largely unburnt by the fire of November 2006 (Table 1). All the tagged banksia seedlings surviving in October 2007 had originated on quadrats sown in March 1991 with seed of their own species, except for three seedlings; a seedling of Banksia aemula on T2D3, another on T3W2 and a seedling of B. oblongifolia on T3W1 (Table 8). All six seedlings of B. aemula that had grown since their lignotubers were first recorded (Table 6) had each originated from seed sown and then shallowly buried (Table 8). In wet heath plot T3W2, the pattern of survival of the relatively numerous seedlings of B. aemula in October 2007 appears to reflect reasonably closely the original 4:6:4 ratio in March 1991 of seed buried: seed sown on disturbed surface: seed sown on undisturbed soil surface among the quadrats on the plot (Table 8). 55 FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS Table 7. Dimensions (mean (S.E.)) of tagged banksia seedlings deemed not to have grown between first recorded presence of lignotuber (in November 1995 unless otherwise indicated) and October 2007. ~ Species NNNBaIsiaacs Ain) ain nD anksiatob oneriol cman Number Initial 2007 Plant | Number Initial 2007 Plant of plants lignotuber lignotuber height of lignotuber lignotuber height width width in plants width width m (mm) (mm) 2007 (mm) (mm) 2007 Dry heath plots T2D2 1 20 33 T2D3 10 15 19 (2) (2) T3D2 9 15 22 (1) (3) Wet heath plots T2W1 5 18 16 (2) (1) T2W3 Die 17 15 T3W1 14 23 DD (2) (2) T3W2 43@ 20 19 11 i 16 15 12 3 14 12 13 (3) (3) 12 ae 12 1] 8 * | plant first record of lignotuber in March1998; @ 4 plants first record of lignotuber in November 1996, and 4 in March 1998. DISCUSSION Fire and habitat interaction Fire and habitat variation interact in different ways across the four species of this study. The interaction is more complex in the two resprouting species than in the two obligate-seeding species. Of the two obligate-seeding species, Dillwynia floribunda has the more straightforward relation with habitat and fire. After fire, seedlings emerge from seeds whose dormancy has been broken by heat, as in Acacia ulicifolia (Auld and O’Connell 1991). Though its seedlings can appear in dry heath, they only survived to maturity in wet heath (Table 3). In wet heath, its soil seed-bank was found by Myerscough et al. (1996) to be abundant, survival of plants after six months was high (Table 3), it was 56 seen to be in flower three and a half years after fire and to be one of the emergent species in the canopy seven and a half years after fire (Table 4). It is one of a suite of obligate-seeding species with soil seed banks and similar sparsely branched erect stems with microphyllous leaves that emerge above resprouting monocotyledons characteristic of wet heath. Other such species are the heaths Epacris microphylla, E. obtusifolia, Sprengelia incarnata, S. sprengelioides, some of which occur with D. floribunda in fire-prone wet heaths on sandstones in the Sydney region (e.g., Keith and Myerscough 1993, Keith 1994, Keith et al. 2007a). Seedlings of Acacia ulicifolia arose in both wet and dry heath particularly after shallow burial of heat-treated seed (Clarke et al. 1996). Survival varied among plots in both wet and dry heath, but there Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH Table 8. Number of tagged banksia seedlings live in October 2007 (grown: plants deemed to have grown; dwarfs: plants deemed not to have grown since their lignotubers were first recorded) in relation to how their seed was placed in March 1991 on or within the soil. Species Banksia aemula Banksia oblongifolia Seed placed Buried Surface Surface not Buried Surface Surface not disturbed disturbed disturbed disturbed Dry heath plots T2D2 dwarf 1 T2D3 dwarfs 2 5° 3 T3D2 dwarfs 1 4 4 grown | Wet heath plots T2W1 dwarfs 1 4 T2W3 dwarfs ] l 1 grown 4 1 1 T3W1 dwarfs 7 6 1 Oh i T3W2 dwarfs 12 I) 12% 2 grown i * includes one seedling that arose in a quadrat not sown with seed of that species. was at least one survivor in each plot immediately before the fire in January 1998 burned six of the plots (Table 3). Some plants were observed to flower at about three and a half years old, and, in an unburnt dry heath plot, plants flowered and set fruit until they were at least seventeen years old. The seedlings are, compared to those of Dillwynia floribunda, slow to gain in height, and are quickly overtopped in wet heath by resprouting monocotyledons, such as Gymnoschoenus sphaerocephalus in T2W1. A modest seed bank of A4.ulicifolia was shown to occur in dry heath but none was found in wet heath (Myerscough et al. 1996). Thus, in wet heath, lack of seed and, for any seed reaching it, scarcity of conditions for successful seedling emergence appear to exclude Acacia ulicifolia, and occurrence of the species is confined to dry heath where it has a soil seed bank, suitable conditions occur after fire for germination of seed and emergence of seedlings, and seedlings are less readily overtopped by other understorey species. Thus Dillwynia floribunda and Acacia ulicifolia are excluded from each other’s characteristic habitat Proc. Linn. Soc. N.S.W., 130, 2009 early in the life cycle, though at different stages; A. ulicifolia through lack of available seed and suitable safe sites (sensu Harper 1977) for any rare seeds present in wet heath, and D. floribunda apparently by lack of suitable growing conditions for seedlings in dry heath. In short, their respective distributions relate to their regeneration niches (sensu Grubb 1977). Beyond the regeneration stage, they need to reproduce successfully in their respective habitats, which observations in this study, while not detailed, indicate occurs, with some seedlings of each species in their fourth year probably contributing seed to the soil seed-bank. This study reveals that in this coastal heath the two resprouting species have three critical phases in their life cycle, regeneration, persistence and growth. What happens to individual plants as they enter and pass through each phase and make transition from one stage to the next depends on fire and habitat. This differs between the two species. Transition from seedling to persistent plant is made evident through fire, while that from fire-resistant but merely persistent plants 57 FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS to plants growing toward reproductive capability is more gradual, presumably depending on success in garnering necessary resources. In the regeneration phase, patterns of seedling establishment between habitats (Myerscough et al. 1996) and among experimental treatments and plots within habitats (Clarke et al. 1996) differed between Banksia aemula and B. oblongifolia. Seedlings of B. oblongifolia that arose in dry heath were fewer and died earlier than on wet heath plots, and, though several survived on T2D3 for six and a half years, none survived the fire in January 1998 (Table 3). In contrast, survival of Banksia aemula occurred across both habitats, and on all the wet heath plots and on three of the dry heath plots there was at least one survivor after seventeen years (Table 3). Survival of seedlings of B. aemula was least on dry heath plots (T2D2 and T3D1) with low water tables (Tables 2 and 3). The fire of 1 January 1998 caused mortality among seedlings of both species, but mortality was much greater in B. oblongifolia than in B. aemula. Overall, in the wet heath plots, 77% of seedlings of B. aemula survived the fire while only 17% did in B. oblongifolia. In both species, to persist through the fire a lignotuber was essential (Table 5). Lignotubers had formed by four and a half years from the sowing of the seed on most of the seedlings that survived, but some had formed somewhat later (Table 7). One factor in the lower survival of seedlings of Banksia oblongifolia is the structure of the lignotubers its seedlings form. They are small and lack the thick corky bark of the larger lignotubers of the seedlings of B. aemula. In B. oblongifolia many of the unburied lignotubers formed completely above the ground surface while this did not occur in seedlings of B. aemula; in them, a lower part was at least in the ground. In both species, as Auld (1987) showed in seedlings of Angophora hispida, burial of the lignotuber enhanced survival of the seedlings, though again to a greater extent in Banksia aemula than in B. oblongifolia. In short, the lignotubers of seedlings of B. aemula appear to be better insulated than those of seedlings of B. oblongifolia, and the transition of seedlings through fire to the fire-resistant persistent phase is made with much less mortality in B. aemula than in B. oblongifolia. In each of the banksia species, very few of the surviving seedlings showed detectable growth between March 1998 and September 2007. Most of them appeared to be simply persisting without detectable net growth. They seem to be in a prolonged “sit-and-wait” state, ageing juvenile plants that Silvertown (1982) called oskars. In many plant communities, growth of 58 such oskars is restricted by lack of sufficient light. Though some shading occurs in the heaths, especially wet heath with abundant monocots in the understorey (Tables 2 and 4), lack of growth in these banksia seedlings is not solely related to light (Table 4). Indeed light was abundant at ground level for several weeks after fire, as occurred when the seedlings arose from seed in 1991 and immediately following their survival through the fire of 1 January 1998. If their growth is resource-limited, the critical resources are those in the soil. Water is probably readily available across the range of habitats, though water stress may be a factor in dry heath with deeper water tables (Table 2). The limiting resources are likely to be one or more of the mineral nutrients needed for plant growth. Previous work (Myerscough and Carolin 1986) has indicated that the sands on which these heaths occur are very low in mineral nutrients. Circumstantial evidence that shortage of mineral nutrients retarded growth of the seedlings comes from the seedlings that grew. All except two were from the wet heath plot T2W3. On this site, when holes were drilled to observe the watertable, a very consolidated coffee rock, B horizon, was reached at c. 0.5 m in three of the four holes. In other wet heath plots, B horizons were deeper and less consolidated. Data of Griffith et al. (2004) indicate that roots of seedlings of both species of banksia, particularly B. aemula, may grow down to B horizons fairly rapidly in similar heaths. It is thus possible that banksia seedlings on this plot could reach the B horizon relatively easily and extract nutrients from it. Furthermore, the plant that grew early and reached reproductive maturity was relatively near one of the holes that had pierced the B horizon to observe the watertable; the disturbance of the hole may have released nutrients that accelerated its growth. Incidentally, this individual flowered when it was less than a metre high (Table 6), showing no sign of requiring an elongated stem for flowering, as in Banksia serrata (Bradstock and Myerscough 1988). Its inflorescences were produced more or less sessile on a thickened main stem that was merely an upward extension of the thickening of the lignotuber. Other low-growing reproductive individuals of B. aemula on this sand system had a similar growth form, while taller growing individuals with trunks occur in sites such as T2D2 and T3D1, and, after fire, produce inflorescences on newly grown stems up to | m long. Growth leading to mature reproductive individuals, persistence of fire-resistant juveniles and regeneration in terms of establishment of seedlings appear as fairly distinct phases in the life cycles of B. aemula and B. oblongifolia. Each phase has its characteristic relations with habitat and fire that differ Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH between the species. In B. oblongifolia, its restriction to wet heath is clearly evident at the regeneration phase (Myerscough et al. 1996), and, as seen in this study, should seedlings survive in a dry heath site they tend to be eliminated in the first fire and thus never enter the phase of fire-resistant juveniles (Tables 3 and 5). Fire-resistant juveniles of B. oblongifolia occurred in wet heath plots, but in the plot, T2W1, having survived the fire of 1 January 1998, they were eliminated (Table 3), probably shaded out under the high cover of Empodisma minus and Gymnoschoenus sphaerocephalus (Table 2). In the other three wet heath plots some continued to survive to seventeen years, but only clearly entering the growth phase in one, T2W3. In Banksia aemula, given availability of seed and modification of the soil surface (Myerscough et al. 1996, Clarke et al. 1996), seedlings arose and survived in all wet and dry heath plots. Ongoing survival was least in the two dry heath plots T2D2 and T3D1 (Table 3) with deep watertables. Transition through the fire of 1 January 1998 on six of the eight plots to persistent fire-resistant juveniles was made with high rates of survival, particularly in the wet heath plots (Table 3). The question arises as to whether the patterns seen at the regeneration stage in seedlings in relation to particular soil treatments applied at sowing of the seeds in March 1991 (see Clarke et al. 1996) were maintained or altered in subsequent survival. In T3W2, the plot with highest number of fire-resistant juveniles persisting at seventeen years, the indication is that the pattern seen in the regeneration phase in relation to soil surface disturbance and seed burial is retained at seventeen years in the persistence phase (Table 8). This plot incidentally was unique among the four wet heath plots in showing little effect of soil treatment and seed burial in numbers of seedlings Surviving at the regeneration phase (see Fig. 2 of Clarke et al. 1996); the three other plots all showed that the greatest number of seedings arose from buried seeds. All six of the plants that were deemed to have entered the growth phase had arisen from buried seed (Table 8). These findings give some insight into the status of populations of the two banksia species on the Eurundereee Pleistocene beach ridges. Firstly, they suggest that their population turn-over is very slow. Indeed, after seventeen years, there is little firm evidence of effective recruitment in either species. In Banksia aemula, only one surviving juvenile showed any evidence of growth in dry heath. While, in wet heath plots, there were numbers of persistent fire- resistant juveniles, a few of which grew, it was an artificial situation brought about by firstly unnaturally Proc. Linn. Soc. N.S.W., 130, 2009 increased availability of seed, relative to naturally occurring levels of seed (Myerscough et al. 1996), and secondly by burial of seeds which is unlikely to occur readily in nature in wet heath (Clarke et al. 1996). In Banksia oblongifolia, very few seedlings survived the fire on 1 January 1998 and persisted as fire-resistant juveniles. The only two that grew arose from seed that in one case had been buried and in the other from seed on a disturbed surface (Table 8). Casual observation of existing mature individuals of either B. aemula or B. oblongifolia suggests that over the seventeen years there was little if any mortality among them. The picture then is of populations of mature long-lived individuals into which there is little opportunity for recruitment of juveniles. Secondly, it appears that, though juveniles may persist several years in a non-growing state, they are limited by lack of resources for growth to progress to mature plants. It is probable that on most plots, the limiting resources are soil nutrients. In the case of one wet heath site with cover of Empodisma minus and Gymnoschoenus sphaerocephalus, lack of light may eliminate juveniles of Banksia oblongifolia, even though mature plants of the species had appreciable cover (Table 2). This suggests that either the current mature plants recruited as seedlings before E. minus and G. sphaerocephalus were so abundant in the site, or, if they were present and abundant, fire frequency was so high that shade from them did not eliminate juveniles of B. oblongifolia. Though the evidence indicates that, presently on these Pleistocene sand ridges, niches for effective regeneration, persistence and growth for these two resprouting species are rare, there must be periods when they are in colonising mode and these niches are more common. This would have been so for B. aemula when parts of Holocene dunes south of Mungo Brush between the Myall River and the sea were colonised by it. Their winged seeds, dispersed some distance in wind, as in those of Banksia serrata observed by Hammill et al. (1998), particularly in willy-willies as were seen to occur in the area of this study on the Pleistocene beach ridges after an intense fire in January 1991, appear well suited for the initial step in colonisation of new habitat. Selection and mode of regeneration The contrast is stark between the two obligate- seeding species studied in which in suitable habitat regeneration is followed very quickly by reproductive maturity of individuals, and the two resprouting species where formation of a fire-resistant lignotuber occurs early and fire-resistant individuals enter a period of persistence in which the majority in this 59 FIRE AND HABITAT INTERACTIONS IN HEATH PLANTS study showed no demonstrable growth toward maturity. As Keith et al. (2007b) have pointed out, the resprouters thus show all the characteristics of Grime’s (1979) stress-tolerators or Stearns’ (1976) K-selected species, while the obligate-seeders are examples of Stearns’ r-selected species. Whether, under selection, their breeding systems follow the suggestion of Heslop-Harrison (1964, Table IV, p. 200) that species with short life cycles, exemplified here by obligate- seeders, are more likely to be inbreeders while species with longer life cycles and slowly maturing adults, exemplified by resprouters, are more likely to be out- breeders would be interesting to establish. It is possible that paths to extinction may differ between obligate-seeders and resprouters. Resprouters may lose effective reproduction through seedlings and reachaterminable state ofa few mature long-persisting individuals, perhaps propagating as clones, while high levels of inbreeding may lead to extinction in some obligate-seeders. How far, in fire-prone habitats, general differences exist between obligate-seeders and resprouters in degrees of in and out-breeding, and thus levels of heterozygosity of individuals, is a question that is yet to be investigated. There is an indication in Table 2 that the species with high cover ten years from fire in both habitats are either strongly obligate-seeding or resprouting, with the possible exception of Pseudanthus orientalis. This would support the suggestion that, in vegetation subject to fairly frequent fires, as appears to have been so in these heaths (Myerscough and Clarke 2007), selection is strong for individuals and thus species to be either markedly obligate-seeding or strongly resprouting and against individuals and species that are neither markedly one nor the other. To establish this as a general rule would require further work. ACKNOWLEDGEMENTS The work began with support from an ARC Small Grant (1990-92) in collaboration with Nicholas Skelton and Peter Clarke. Beside Nicholas Skelton and Peter Clarke, Neil Tridgell, Ian Radford, Joan Myerscough, Andrew Denham, Alan Keating and James Myerscough each helped in the field, especially Neil Tridgell who accompanied me many times; I thank them all. I thank Tony Auld for the fire-proof tags used with the banksia seedlings, and Tony Auld and Andrew Denham for loan of callipers modified by Murray Ellis for measuring diameters of lignotubers. The work was done under licence from the Director of the New South Wales National Parks and Wildlife Service. Staff of Myall Lakes National Park helped with access to study sites, which is much appreciated. I thank an anonymous referee for constructive comments, and Jan Percival for help in getting the paper into correct electronic form for publishing. 60 REFERENCES Auld, T.D. (1987). Post-fire demography in the resprouting shrub Angophora hispida (Sm.) Blaxell: flowering, seed production, dispersal, seedling establishment and survival. Proceedings of the Linnean Society of New South Wales 109, 259-269. Auld, T.D. and O’Connell, M.A. (1991). Predicting patterns of post-fire germination in 35 eastern Australian Fabaceae. Australian Journal of Ecology 16, 53-70. Beadle, N.C.W. (1940). Soil temperatures during forest fires and their effect on the survival of vegetation. Journal of Ecology 28, 180-192. Beadle, N.C.W. (1981). “The vegetation of Australia’. (Cambridge University Press, Cambridge). Benwell, A.S. (1998). Post-fire recruitment in coastal heathland in relation to regeneration strategy and habitat. Australian Journal of Botany 46, 75-101. Bond, W.J. and Midgley, J.J. (2001). Ecology of sprouting in woody plants: the persistence niche. 7rends in Ecology and Evolution 16, 45-51. Bradstock, R.A. and Myerscough, P.J. (1988). The survival and population response to frequent fires of two woody resprouters Banksia serrata and Isopogon anemonifolius. Australian Journal of Botany 36, 415- 431. Carolin, R.C. (1970). Myall Lakes - an ancient and modern monument. The Proceedings of the Ecological Society of Australia 5, 123-129. Clarke, P.J., Myerscough, P.J. and Skelton, N.J. (1996). Plant coexistence in coastal heaths: between- and within-habitat effects of competition, disturbance and predation in the post-fire environment. Australian Journal of Ecology 21, 55-63. Clifford, H.T. and Specht, R.L. (1979). ‘The vegetation of North Stradbroke Island’. (University of Queensland Press, St Lucia, Queensland). Gill, A.M. (1981). Coping with fire. In “The biology of Australian plants’ (Eds. J.S. Pate and A.J. Mc Comb) pp. 65-87. (University of Western Australia Press, Nedlands). Griffith, S.J., Bale, C., Adam, P. and Wilson R. (2003). Wallum and related vegetation on the NSW North Coast: description and phytosociological studies. Cunninghamia 8, 202-252. Griffith, S.J., Bale, C. and Adam, P. (2004). The influence of fire and rainfall upon seedling recruitment in sand- mass (wallum) heathland of north-eastern New South Wales. Australian Journal of Botany 52, 93-118. Grime, J.P. (1979). ‘Plant strategies and vegetation processes’. (J. Wiley and Sons: London). Grubb, P.J. (1977). The maintenance of species richness in plant communities: the importance of the regeneration niche. Biological Reviews 52, 107-145. Hammill, K.A., Bradstock, R.A. and Allaway, W.G. (1998). Post-fire fire seed dispersal and species re- establishment in proteacous heath. Australian Journal of Botany 46, 407-419. Proc. Linn. Soc. N.S.W., 130, 2009 P.J. MYERSCOUGH Harden, G.J. (1990). “Flora of New South Wales, Volume 1°. (University of New South Wales Press, Sydney). Harden G.J. (1992). ‘Flora of New South Wales, Volume 3’. (University of New South Wales Press, Sydney). Harden, G.J. (1993). “Flora of New South Wales, Volume 4’. (University of New South Wales Press, Sydney). Harden, G.J. (2002). ‘Flora of New South Wales, Volume 2° (revised edition). (University of New South Wales Press, Sydney). Harper, J.L. (1977). ‘Population biology of plants’. (Academic Press: London). Heslop-Harrison, J. (1964). Forty years of genecology. Advances in Ecological Research 2, 159-247. Keith, D.A. (1994). Floristics, structure and diversity of natural vegetation in the O’Hares Creek catchment, south of Sydney. Cunninghamia 3, 543-594. Keith, D.A. (2004). “Ocean shores to desert dunes: the native vegetation of New South Wales and the ACT’. (Department of Environment and Conservation (NSW), Hurstville). Keith, D.A. and Myerscough, P.J. (1993). Floristics and soil relations of upland swamp vegetation near Sydney. Australian Journal of Ecology 18, 325-344. Keith, D.A., Holman, L., Rodoreda, S., Lemmon, J. and Bedward, M. (2007a). Plant functional types can predict decade-scale changes in fire-prone vegetation. Journal of Ecology 95, 1324-1337. 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Seedling growth and storage characterisitics of seeder and resprouter species of Mediterranean-type ecosystems of S.W. Australia. Annals of Botany 65, 585-601. Silvertown, J.W. (1982). ‘Introduction to plant population ecology’. (Longman, London and New York). Specht, R.L. (1981). The sclerophyllous (heath) vegetation of Australia: the eastern and central states. In ‘Ecosystems of the world. Vol 9A. Heathlands snd related shrublands. Descriptive studies’ (Ed. R.L. Specht) pp. 125-210. (Elsevier, Amsterdam). Proc. Linn. Soc. N.S.W., 130, 2009 Stearns, S.C. (1976). Life-history tactics: a review of ideas. The Quarterly Review of Biology 51, 3-47. Thom, B.G., Shepherd, M., Ly, C.K., Roy, P.S., Bowman, G.M. and Hesp, P.A. (1992). Coastal geomorphology and Quaternary geology of the Port Stephens-Myall Lakes Area. Department of Biogeography and Geomorphology, ANU Monograph No. 6, Australian National University, Canberra. 61 FURY AND HLA BIT ARHDUDDE RAINE — PL pee bbs At Hert. \gatoenar nity vane MT COB A ab Mi asad iN: dausytt F964, we AS cepts, tA oF. wed) Yr ‘eRe o gral ied) | bare dg’ aie \ reratagt! wy ii i ‘yale Neg tv) ~wrtaley ee a | iene rts rah Bas ” nsehyhO WPT ARIE “APA NEAR pd S58 1 cme ites 4 pha lte ‘fond Li sa iid lil oindains sl rey, riots ney hansen PL ia WA). e ie 7 Thycanty t OME hire PPLE Ral RIS iy area FS CinePinnts any waite parte Tyee: oy Te mime te ay hry nny a pe ANE, by iy S hele Gath ey Re a ete eer ¢ asl Nevl A eedig tae Ip \ of] fae I el a) Vay mind |) Fix +, Dt ag ie Galt rk eat Why me ae rp TANK, rt hat ‘thw He euiny Sirgen hd tas iTiee ‘ i ie tian TOR, Fae vw Koel teil mee ei, Oh nti eine syetpoxe fiw ee iva! jaty #9 ae Py w4 real BS! ct an) At ‘Tey Het les ir ey We + a ‘ ay n Tie aig Jt iS abe HF He) ‘am phe nevis W? Chars o7 ed weeadtes a “ABS DMA: SID. 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NOP re oO epee RIM, og Civile, SILT IEE Hidlpet Rigeh amok gis neightih bef etek or RortaaAP chery ts cour wha HANNS / held Wah e iis beth ee Tele Bite Ay SE SPARS) Site SPA SS tlh papraray OPES aa ih i Late Llandovery (Early Silurian) Dendroid Graptolites from the Cotton Formation near Forbes, New South Wales R. B. Rickarps!, A. J. WRIGHT? AND G. THOMAS? 'Department of Earth Sciences, University of Cambridge, Downing Street, Cambridge CB2 3EQ, U.K. (rbr1000@esc.cam.ac.uk); *School of Earth and Environmental Sciences, University of Wollongong, Wollongong NSW 2522 (tony_ wright@uow.edu.au) and Linnean Macleay Fellow; 3P.O. Box 130, Southland Centre, Victoria 3192. Rickards, R.B., Wright, A.J. and Thomas, G. (2009). Late Llandovery (Early Silurian) dendroid graptolites from the Cotton Formation near Forbes, New South Wales. Proceedings of the Linnean Society of New South Wales 130, 63-76. A well-preserved dendroid graptolite fauna of Early Silurian (late Llandovery: probable turriculatus graptolite zone) age is described from the Cotton Formation near Forbes, New South Wales. A possible rhabdopleuran hemichordate is described from Australia for the first time. The fauna consists of 13 taxa as follows: Dendrograptus sp. aff. D. avonleaensis, Dictyonema zalasiewiczi sp. noy., Dictyonema sp. aff. D. paululum australis, Dictyonema paululum australis, Dictyonema sp. aff. D. sp. cf. D. venustus of Bulman (?ssp. nov.), Dictyonema venustum, Dictyonema sp. ef. D. falciferum, Callograptus bridgecreekensis, Callograptus rigbyae, Callograptus sp. aff. C. ulahensis, Stelechocladia sp. cf. S. praeattenuata, Acanthograptus praedeckeri and ?Rhabdopleura sp. (? with zooids). The fauna is close in composition (although less diverse) and age to a dendroid fauna recently described from Bridge Creek near Orange, NSW, which was assigned to the slightly younger griestoniensis zone. Manuscript received 12 March 2008, accepted for publication 23 January 2009. KEY WORDS: Cotton Formation, dendroids, Early Silurian, Forbes, graptolites, New South Wales. INTRODUCTION The dendroid graptolites described here have been collected over many years by one of us (GT) from a quarry in the Cotton Formation at Cotton Hill near Forbes in western N.S.W. Fossils from these beds have been described by Sherwin (1974: graptolites) and Edgecombe and Sherwin (2001: trilobites). The described trilobite and graptolite faunas are from beds exposed in the quarry high in the upper part of the Cotton Formation (Sherwin 1974) and the graptolite fauna is correlated with the late Llandovery (Early Silurian) turriculatus graptolite zone (Edgecombe and Sherwin 2001). Despite the very nature of collections made in an active quarry, there seems little doubt that the bulk of the dendroid fauna and the graptoloids are from the same narrow horizon. The most similar known dendroid fauna was described by Rickards et al. (2003) from the Four Mile Creek district, south of Orange, NSW, and comparisons are made below with that fauna. AGE OF THE ASSEMBLAGE Although the Cotton Formation dendroid fauna (13 species and subspecies) is less diverse that described from the Bridge Creek localities in the Four Mile Creek district (24 species and subspecies) by Rickards et al. (2003), there can be little doubt that the two faunas are close in age. The largest assemblage at Bridge Creek, from locality F14, was referred by Rickards et al. (2003) to a horizon low in the griestoniensis graptolite Zone. The Cotton Hill fauna is assigned almost certainly to the stratigraphically lower turriculatus graptolite zone. Of the fauna we record here from the Cotton Formation, only Dictyonema zalasiewiczi sp. nov. and ?Rhabdopleura sp. have not been recorded from Bridge Creek at locality F1l4. Callograptus ulahensis Rickards et al., 2003 was recorded from a lower (gregarius Zone) assemblage at locality BF15 on Bridge Creek: the Cotton Hill Quarry species is referred to Callograptus sp. cf. C. ulahensis. EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES Stelechocladia praeattenuata Rickards et al., 2003 was not recorded from F14 but occurs below (F19) and above (BF28, BF24 and BF 18), ranging from the gregarius Zone to the uppermost griestoniensis Zone. Sherwin (1973, 1974) referred the strata at Cotton Hill Quarry to the turriculatus Zone, with some levels probably earlier than this but without definite faunas. Sherwin (1970, 1973) also recorded Dictyonema spp. from the highest band of a group of beds yielding a likely turriculatus Zone fauna. Hence the two dendroid assemblages, from Cotton Hill (probable turriculatus Zone) and from Bridge Creek (griestoniensis Zone), are not dissimilar in age, the Cotton Hill fauna being about one graptolite zone lower. There is another difference between the two assemblages apart from a possible slight age difference and a diversity range, and that is that the Cotton Hill fauna is almost exclusively of slender, delicate species, often broken. In contrast, most of the species described by Rickards et al. (2003) from Four Mile Creek are robust, and are preserved in poorly bedded siltstone. The only robust form common to the two localities is Stelechocladia and at Cotton Hill it is known only from three small fragments showing distal, slender thecae. It is possible that the Cotton Hill assemblage lived in a quieter depositional environment, such as a lagoon, or further offshore. Edgecombe and Sherwin (2001) concluded that the laminated siltstones that dominate the formation were deposited in a ‘very calm’ environment, ‘most likely below storm surge wave base’. Associated graptoloids. Sherwin(1970, 1973) was the first to identify graptoloid species from the Cotton Beds, following the initial recognition of graptolites from this locality by Packham (1967). Sherwin (1970, 1973) recognised two faunas, an earlier assemblage (his fauna C) and a later assemblage (his fauna D) respectively from the east and west quarries on Cotton Hill: both are in the upper Cotton Formation. Fauna D, from the western, larger quarry, includes Dictyonema sp. (Sherwin 1973, fig. 10). Some mixing of faunas possibly occurred because collection was from large blocks on the quarry floor (Sherwin 1974, p. 149). It is this western quarry from which the present collection of dendroids came; the eastern, smaller, quarry has not so far yielded dendroid graptolites. The graptoloid assemblages were described in detail by Sherwin (1974) and, allowing for some possible mixing of faunas, the overall aspect is of a turriculatus Zone fauna, perhaps rather low in that horizon given the presence of Rastrites linnaei, Monograptus halli, and Monograptus sp. cf. M. sedgwickii. Thus the Cotton Hill quarry is at 64 a stratigraphically lower level than the Four Mile Creek (F14) locality which was mentioned in the preceding section and which is probably low in the griestoniensis Zone. Graptoloids occurring on the same rocks as the Cotton Hill dendroids described below include: Parapetalolithuspalmeus, ? Glyptograptus tamariscus, Monograptus andrewsi and Spirograptus turriculatus (Fig. 7b). The faunal lists given by Sherwin (1974) are fuller and much more reliable than the graptoloids at our disposal. Here we also record and illustrate (Fig. 7a) Parapetalothius palmeus (Barrande, 1850), a form not recorded by Sherwin (1974); its occurrence accords with his age attribution of the turriculatus zone. In their revision of Spirograptus, Loydell et al. (1993) assigned Sherwin’s (1974) Monograptus turriculatus (Barrande, 1850) to their new species Spirograptus guerichi. They further (Loydell et al. 1993, p. 924, text-fig. 7) stated that S. guerichi is “virtually confined to its biozone”, whereas S. turriculatus ranges through their turriculatus biozone into the crispus biozone; Sherwin (1974, p. 150) shows that both species occur in his fauna D. SYSTEMATIC PALAEONTOLOGY This benthic graptolite fauna has been assembled only by sustained and diligent collecting over many years by one of us (GT), as dendroids are rare at the locality. The preservation is of reddish brown graptolites against a very pale, fine-grained siltstone or mudstone. The specimens are often large but are mostly fragmentary, and there seems to be little in the way of burial distortion or twisting and no obvious tectonic deformation. Some specimens are preserved in three dimensions infilled, probably with goethite: in others the periderm is diagenetically flattened, but with some parts (e.g. stolons) pyritised. It is possible that in some instances pyritised zooids are present. Rarely stolons occur free on the bedding plane, the surrounding periderm having degenerated; this situation has been noted by Chapman et al. (1993) and Rickards et al. (2003). All specimens are deposited in the Australian Museum, Sydney, with numbers AM F123381-123428. Subphylum Pterobranchia Lankester, 1877 (nom. trans. Rickards and Durman 2006) Class Graptolithina Bronn, 1849 Order Dendroidea Nicholson, 1872 Family Dendrograptidae Roemer in Frech 1897 Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Dendrograptus J. Hall, 1858 Synonymy aff. 2003 Dendrograptus avonleaensis n. sp.; Type species Rickards et al., pp. 312-3, figs 5A, 6A. Graptolithus hallianus Prout, 1851, subsequently designated by J. Hall (1862). Material AM F123381. Dendrograptus sp. aff. D. avonleaensis Rickards et al., 2003 Description Figures la, 3a The single specimen shows nine stipes and five Figure 1. a, Dendrograptus sp. aff. D. avonleaensis Rickards et al., 2003; AM F123381. b, Dictyonema sp. aff. D. cf. venustum Bulman, 1928; AM F123398. c, Dictyonema zalasiewiczi sp. nov., holotype AM F123402; d, Callograptus bridgecreekensis Rickards et al., 2003; AM F123403. e, Callograptus rigbyae Rickards et al., 2003; AM F123406. Black bars are dissepiments; black rods, arched in ventral views, are autothecal ventral processes. Scale bars 1 mm. Proc. Linn. Soc. N.S.W., 130, 2009 65 EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES Figure 2. a-b, Dictyonema paululum australis Rickards et al., 2003, respectively AM F123382, AM F123383. c-d, Acanthograptus praedeckeri Rickards et al., 2003, respectively AM F123411, AM F123410. Scale bars 1 mm. branching points in mostly ventral view, but also shows autothecal profiles in places. Specimen almost flattened diagenetically, but periderm still slightly transparent. 15-20 autothecae in 10 mm, with a profile width of 0.2 mm, and a fairly simple aperture slightly arched ventrally as seen in the ventral view. Bithecae exceedingly inconspicuous, being tiny tubes opening externally alongside autothecal apertures and alternating along stipe. Branching of stipes may be in zones at 1-3 mm intervals; stipe lateral width 0.30 mm and dorsoventral width 0.40 mm. Remarks This specimen, probably representing the distal- 66 most parts of the colony, agrees closely with the type material of D. avonleaensis in most characters, especially the roughly zonal branching, autothecal nature and spacing and stipe dimensions. The type specimens from Bridge Creek (Rickards et al. 2003) had much of the proximal region preserved, and this is much more robust than the Cotton Hill material. Dictyonema J. Hall, 1851 Type species Gorgonia retiformis J. Hall, 1843, subsequently designated by Miller (1889). Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Figure 3. a, Dendrograptus sp. aff. D. avonleaensis Rickards et al., 2003; AM F123381. b, Callograptus bridgecreekensis Rickards et al., 2003; AM F123403. c, Callograptus sp. aff. C. ulahensis Rickards et al., 2003; AM F123407. d, Dictyonema sp. cf. D. falciferum Bulman, 1928, AM F 123401. e, Dictyonema paululum australis Rickards et al., 2003; AM F123386. Scale bars 1 mm. Proc. Linn. Soc. N.S.W., 130, 2009 67 EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES Figure 4. a, Dictyonema zalasiewiczi, sp. nov., holotype AM F123402. b, Dictyonema venustum Lapworth, 1881; AM F123397. c, Dictyonema paululum australis Rickards et al., 2003, respectively AM F123385. Scale bars 1 mm. Dictyonema paululum australis Rickards et al., 2003 Figures 2a-b, 3e, 4c, 7c Synonymy 2003 Dictyonema paululum australis n. subsp.., Rickards et al., p. 316, figs 7F-G, 9E, 12A. Material Twelve specimens, AM F123382-93, ranging from small fragments to almost complete colonies. 68 Description Probably fan-shaped rhabdosome of slender stipes, no indication of a conical colonial arrangement; colony with slender, parallel stipes, only approximately branching in zones, sometimes fanning out in rapid expansion. Stipes branch at intervals of 1.5-3.0 mm; stipe lateral width 0.20-0.25 mm proximally, 0.15 mm more distally; dorsoventral width 0.50-0.60 mm; stipe spacing 13-16 in 10 mm, stipe interspaces 0.50-0.60 mm. Autothecae denticulate, 18-20 in 10 Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Figure 5. a, Dictyonema sp. cf. D. falciferum Bulman, 1928, partially preserved stolons in three portions of stipes where periderm is degenerate; AM F123400. b, Stelechocladia sp. cf. S. praeattenuata Rick- ards et al., 2003, AM F123408a. c, Acanthograptus praedeckeri Rickards et al., 2003, AM F123410. d-e, ?Rhabdopleura sp., respectively AM F123412-3; both exhibit stolons with possible preserved soft tissue (encysted zooidal attached). f, Callograptus sp. aff. C. ulahensis Rickards et al., 2003; AM F123407. Scale bars 1mm; stipple on Fig. a indicates possible attached soft parts. Scale bars 25 mm (a), 1 mm (b-f). mm; dissepiments slender, 0.05-0.10 mm, 14-20 in 10 mm. Dissepiments conspicuous because of their frequency; proximally they are more robust and perhaps sparser. Bithecal tubes seen in places but their apertural regions are difficult to discern; they may be of the type described by Bulman (1928) in D. falciferum where the bithecal apertural region hooks over the dorsal apertural region of the autotheca. Alternatively, they may grow short of the full hook (Fig. 3e); bithecal tubes 0.05 mm wide. Proc. Linn. Soc. N.S.W., 130, 2009 Remarks Rickards et al. (2003) considered the original material from Four Mile Creek probably had conical rhabdosomes but it seems more likely that they are fan-shaped. Bulman (1928) could not see the nature of the rhabdosome as a whole in the type subspecies, and he was particularly vague about the nature of the bithecae: otherwise the type subspecies is clearly close to the Australian form differing only as outlined by Rickards et al. (2003). Dictvonema paululum australis is the most common dendroid at Cotton Hill Quarry 69 EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES Figure 6. a-b, Dictyonema sp. aff. D. paululum australis Rickards et al., 2003; respectively AM F123395, AM F123396. c, Callograptus rigbyae Rickards et al., 2003; AM F123405. d, Dictyonema sp. aff. D. sp. cf. D. venustum Bulman, 1928; AM F123398. Scale bars 1 mm. 70 Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Figure 7. a, Parapetalolithograptus palmeus (Barrande, 1850) s.1., AM F123428. b, Spirograptus turricula- tus (Barrande, 1850), AM F 123427. c, Dictyonema paululum australis Rickards et al., 2003; AM F123384. d-e, Callograptus rigbyae Rickards et al., 2003; respectively AM F123404, AM F 123406. f, Acanthograp- tus praedeckeri, AM F123409. Scale bars 1 mm. Proc. Linn. Soc. N.S.W., 130, 2009 ral EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES (and see also D. sp. aff. D. p. australis described below). Dictyonema paululum hanoverense Rickards et al., 2005 from the Late Silurian parultimus Zone near Neurea, N.S.W. differs in having an autothecal spacing of 28-30 in 10 mm and quite spinose ventral apertures. Dictyonema sp. aff. D. paululum australis Rickards et al., 2003 Figure 6a-b Synonymy aff. 2003 Dictyonema paululum australis subsp. nov.; Rickards et al., p. 316, figs 7F-G, 9E, 12A. Material AM F123394-6, 123415a-b. Description Nature of colony uncertain, possibly fan-shaped. Stipes with lateral width of 0.20-0.25 mm, and spaced at 20-22 in 10 mm, more or less parallel, and with interstipe spaces of 0.20-0.40 mm; branching roughly in zones every 1.0-2.5 mm. Autothecae spaced at 19- 20 in 10 mm; dorsoventral width uncertain but may be ca. 0.50 mm. Bithecae not detected. Dissepiments fine, spaced at ca. 20 in 10 mm. Remarks These specimens are superficially similar to those of the D. paululum australis material described in this paper, except that the stipes are more closely spaced and the interstipe spaces concomitantly narrow. There may be a temporal subspeciation factor involved here as the source level in the quarry for the specimens is uncertain; thus some of the D. p. australis specimens may be from older beds and others from the turriculatus level. Dictyonema venustum Lapworth, 1881 Figure 4b Synonymy 1881 Dictyonema venustum sp. nov.; Lapworth, pp. 171-2, pl. 7, fig. la-c 1928 Dictyonema venustum, Lapworth, emend; Bulman, pp. 61-3, pl. 5, figs 6-7, ?8, text-fig 34. 2003 Dictyonema venustum Lapworth, 1881; Rickards et al., pp. 315-6, figs 7A, 9D, 10B-D. Material An almost complete rhabdosome, AM F 123397, plus AM F123416-7. 2 Description Rhabdosome conical, reaching 8 mm x 8 mm; very proximal end missing though part of the holdfast may be present. Stipes with lateral width of 0.25-0.30 mm, dorsoventral width of 0.70 mm, and spaced at 16 in 10 mm. Interstipe spaces rectangular, up to 0.50 mm wide, and are bounded by stipes and dissepiments spaced at 5-8 in 10 mm. Dissepiments relatively robust, up to 0.15 mm thick. Autothecal spacing 16 in 10 mm; thecae appear to be denticulate but otherwise simple. Bithecal tubes present but relationships to autothecal apertures not seen. Remarks The specimen is very close to the type material redefined by Bulman (1928), differing only in a slightly closer spacing of the stipes. Dictyonema sp. aff. D. sp. cf. venustum Bulman, 1928 Figures 1b, 6d Synonymy aff. 1928. Dictyonema cf. venustum Lapworth, emend.; Bulman, pp. 62-3, pl. 5, fig. 8 (non 6-7). Material AM F123398; three other specimens (AM F123424-6) questionably assigned here. Description The large fragmental rhabdosome (AM F 123398) has 12 stipes preserved, spaced at 16 in 10 mm, with interstipe spaces of 0.10-0.40 mm, and spaced at ca. 1-6 in 10 mm. Lateral stipe width 0.25 -0.40 mm, usually nearer the latter. Autothecae unclear but may be spaced at ca. 20 in 10 mm with dorsoventral width of 0.50 mm. Remarks This specimen is very close to that figured by Bulman (1928, pl. 5, fig. 8) which he listed as D. venustum but he made it clear in the text that he placed it there only with reserve. As in the Cotton Hill quarry specimen the interstipe spacing 1s less and the stipes are more robust. The Girvan specimens illustrated by Bulman were said to come from communis zone beds (probably convolutus-sedgwickii zone in modern terminology); thus they may have come from pre- turriculatus Zone strata, and this is also possible in the case of the present specimen. Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Dictyonema sp. cf. D. falciferum Bulman, 1928 Figures 3d, 5a Synonymy cf. 1928 Dictyonema falciferum n. sp.; Bulman, pp. 53-6, pl. 5, figs 1-3, text-figs 27-29. cf. 2003 Dictyonema falciferum Bulman, 1928; Rickards et al., p. 315, figs 5I, 8B, 9C, 10A. Material AM F123399-123401. Description Rhabdosome possibly fan-shaped (?conical), at least 25mm long and 18 mm broad, with numerous parallel stipes spaced at 14 in 10 mm, having stipe interspaces of 0.50-0.60 mm. Rectangular meshes are defined by stipes and conspicuous dissepiments spaced at 8-10 in 10 mm. Autothecal spacing 20 in 10 mm. Lateral stipe width 0.20-0.25 mm, and dorsoventral stipe width 0.50 mm. Autothecae appear to be simple denticulate but not spinose. Bithecal tubes present but their apertural regions unclear. Branching rather irregular, at 0.5-5.0 mm intervals. Remarks These specimens are closely similar to the specimens described from Four Mile Creek by Rickards et al. (2003) differing only in having a less regular branching pattern and slightly more parallel stipes. One specimen (AM F123400; Fig. 5a) has traces of preserved stolons. Dictyonema zalasiewiczi sp. nov. Figures Ic, 4a Material Holotype, AM F123402, an almost complete rhabdosome. Derivation of name After Dr. J. Zalasiewicz, University of Leicester, a leading graptolite worker. Diagnosis A Dictyonema species with 30-40 dissepiments in10 mm; stipes 0.2-0.5 mm wide and spaced at 0.2- 0.3 mm. Description Fan-shaped rhabdosome more than 30 mm long and over 20 mm wide, typified by its striking number of dissepiments, up to 40 per 10 mm, never less than 30. Dissepiments 0.05-0.10 mm across, often arched Proc. Linn. Soc. N.S.W., 130, 2009 distally, and quite frequently branching; commonly angled rather than normal to adjacent stipes, but also occur as closely spaced pairs. Stipes uniformly 0.20- 0.25 mm in lateral width, with branching every 2-2.5 mm proximally and more sparse distally, up to 6 mm. Branching occurs in broad zones. Stipes parallel and closely spaced, with interstipe spaces of 0.20-0.30 mm, similar to the lateral width, resulting in a stipe spacing of about 20 in 10 mm. Autothecal spacing difficult to discern in this dorsoventral view, but may be around 20 in 10 mm. Nature of autothecal apertures cannot be seen, except in one area where they appear to be denticulate or spinose. Bithecae not detected. Remarks This is a highly unusual and distinctive species because of the huge number of dissepiments. Bulman (1928, table II) gave only two species of Silurian dictyonemids with as many as 20 dissepiments in 10 mm (and none with this frequency in the Ordovician species; Bulman 1928, table I). Of Australian dictyonemids, Rickards and Wright (1997) and Rickards et al. (2003), for example, only once have dissepimental spacings as high as 30 in 10 mm been recorded, and that in some specimens of Dictyonema delicatulum barnbyensis from the middle to upper Ludlow; a few other Australian species have as many as 20 in 10 mm. Dictyonema paululum australis Rickards et al., 2003 is similar in having conspicuous dissepiments, but their spacing and that of the stipes is quite different. None of Boucek’s (1957) dictyonemids has high dissepimental spacings. Callograptus J. Hall, 1865 Type species Callograptus elegans J. Hall, 1865, by original designation. Callograptus bridgecreekensis Rickards et al., 2003 Figures 1d, 3b Synonymy 2003 Callograptus bridgecreekensis n. sp.; Rickards et al., p. 319, figs 14A, 15A-B. Material AM F123403. Description These 13 or so stipes are towards the distal end of a moderately-sized (8 mm x 5 mm) piece of rhabdosome; lateral stipe width of 0.50 mm most 13 EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES proximally, and 0.20 mm at distal ends of stipes. Branching irregular, stipe spacing over 20 in 10 mm. No dissepiments. Autothecae not detected in this wholly dorsoventral view, but traces of bithecal tubes apparent. Callograptus rigbyae Rickards et al., 2003 Figures le, 6c, 7d-e Synonymy 2003 Callograptus rigbyae n. sp. Rickards et al., p. 319, figs 14B-C. Material Four almost complete colonies, AM F123404-6, 123414, plus AM F123419-123421. Description Fan-shaped or discoidal colony about 10 mm across, developed from a small holdfast. Up to 6 branching zones may occur in this short distance giving numerous peripheral stipes. Rare anastomosis of stipes. Interstipe spacing 0.50 mm; stipe spacing ca. 16 in 10 mm, lateral stipe width 0.20-0.30 mm. Autothecae spaced at 20 in 10 mm, and autothecal apertures bear a ventral spine up to 0.50 mm long. Dissepiments rare, and extremely fine. Bithecae occur, but their nature is unclear. Remarks The original specimens from Bridge Creek (Rickards et al. 2003, p. 319) were two colonies preserved in plan view. Two Cotton Hill specimens (Figs 7d-e) are more in profile. One (AM F123404a- b: Fig. 7d) shows the autothecae best and a short spine can be clearly seen. Bithecae were not detected in the original material. Callograptus sp. aff. C. ulahensis Rickards et al., 2003 Figures 3c, 5f Synonymy aff. 2003 Callograptus ulahensis n. sp.; Rickards et al., pp. 319-20, figs 16A, 17A. Material A small fragment of rhabdosome, AM F 123407, comprising nine stipes. Description The initial two parallel stipes branch after 3 mm, but thereafter branch at 1-1.5 mm intervals resulting in short, parallel stipes with lateral width 74 of 0.20 mm. Interstipe spaces ca. 0.50 mm, and stipe spacing ca. 20 in 10 mm. Autothecal spacing 20 in 10 mm; dorsoventral width may be 0.40-0.50 mm and thecal aperture may be denticulate. No dissepiments present. Remarks This specimen adds a little to the original description which was based upon two specimens (AM F114760 and 114780) from locality BF15, some 100 m S of the junction of Four Mile Creek and its tributary Bridge Creek (Rickards et al. 2003). The autothecae are not so clear in the Cotton Hill Quarry specimen, but the disposition of the stipes is more apparent. Family Stelechocladiidae Chapman et al., 1993 Stelechocladia Pocta, 1894 Type species — Stelechocladia subfruticosa Poéta, 1894, subsequently designated by Boucek (1957). Stelechocladia sp. cf. S. praeattenuata Rickards et al., 2003 Figure 5b Synonymy cf. 2003 Stelechocladia praeattenuata Nn. sp.; Rickards et al., p. 322, figs 17B, 19A-B. Material AM F123408a-b, and AM F123422-3. Description AM F 123408 is the distal end of a stelechocladiid with stipes spaced at 16 in 10 mm, some apparently laterally derived from nearby dominant stipes. Lateral stipe width from 0.20-0.40 mm, the more robust stipes being more proximal. Branching, where it occurs, is almost every mm, but long, unbranched portions also occur. Autothecae not seen. Remarks This form is almost certainly referable to S. praeattenuata, having the typical combination of dichotomous and “lateral” branching as well as the dimension of a distal part of that species’ rhabdosome. Lack of autothecal presentation, however, urges caution. Proc. Linn. Soc. N.S.W., 130, 2009 R.B. RICKARDS, A.J. WRIGHT AND G. THOMAS Family Acanthograptidae Bulman, 1938 Acanthograptus Spencer, 1878 Type species Acanthograptus granti Spencer, 1878, by original designation. Acanthograptus praedeckeri Rickards et al., 2003 Figures 2c-d, 5c, 7f Synonymy 2003 Acanthograptus praedeckeri n. sp.; Rickards et al., pp. 322-5, figs 17C-D, 19C, 20A- (not fig. 18A). 2003 Dictyonema warrisi; Rickards et al., fig. 18A (mislabelled). Material Three specimens, including one almost entire, small rhabdosome (AM F 123409, Fig. 7f): AM F123409-11. Description Twigs arranged at 8-16 in 10 mm, each 0.70- 1.00 mm long and comprising two or more thecae. Main stipes 0.40-0.50 mm wide laterally, and their ramifications fill all the space available to form a flabellate or fan-shaped colony. Branching occurs every 0.50-2.0 mm, usually 1.00-1.50 mm. Autothecal tubes 0.10 mm wide and do not seem to expand towards apertures. Bithecae may be not much smaller and may open near bases of twigs or on main stipe. Remarks The caption for Rickards et al. (2003, fig. 18A) wrongly states that the illustrated species is Dictyonema warrisi, really being Acanthograptus praedeckeri. Class Rhabdopleurina Fowler, 1892 Family Rhadopleuridae Harmer, 1905 Rhabdopleura Allman, 1869 Type species R. normani Allman, 1869. ?Rhabdopleura sp. Figures 5d-e Material AM F1123412-3; the latter has a fragment of Callograptus rigbyae on the reverse side (AM F123414). Proc. Linn. Soc. N.S.W., 130, 2009 Description The larger specimen (AM F123412: Fig. 5d) appears to have a basal thecorhiza from which arise about nine tubes with a diameter of 0.15-0.20 mm. Tubes distally less sclerotised. Suggestion of growth lines in places, especially on AM F123413 (Fig. 5e). In thecorhizal portion there are probably pyritised (non-goethitised) stolons and possibly also attached encysted zooids. Distal parts of tubes (coenecia) unoccupied and may represent free-standing parts of tubes. AM F123413 may also have pyritised stolons and zooidal remains. Remarks This form does not resemble tuboids such as Galeograptus and Cyclograptus which we have previously recorded from Australia (Rickards et al. 1995, 2003). Were it not for the uncertainty about the growth lines we would refer this to Rhabdopleura with more confidence. Rhabdopleura has not previously been recorded in Australian strata. ACKNOWLEDGEMENTS. RBR acknowledges support from Emmanuel College; the Department of Earth Sciences, University of Cambridge; and the Royal Society of London. AJW acknowledges with much gratitude the award of the Linnean Macleay Fellowship during 2006-7-8 by the Linnean Society of New South Wales, and the support of the School of Earth and Environmental Sciences, University of Wollongong. We are particularly grateful to Dudley Simon (Department of Earth Sciences, Cambridge University) for his skilful assistance in taking all the photographs and processing them. We also thank the reviewers for comments which considerably improved the original manuscript. REFERENCES Allman, G.T. (1869). On Rhabdopleura, a new genus of Polyzoa. Proceedings of the Royal Society of Edinburgh 6, 438-440. Barrande, J. (1850). Graptolites de Bohéme. Extrait du Systéme Silurien de la Bohéme. 74 pp. Published by the author, Prague. Bouéek, B. (1957). The dendroid graptolites of the Silurian of Bohemia. Sbornik Ustredniho Ustavu Geologického 23, 294 pp, Prague. Bronn, H.G. (1849). Index Palaeontologicus B, Enumerator Palaeontologicus. 980 pp. Stuttgart, E., Schweizerbart’sche. TD EARLY SILURIAN GRAPTOLITES FROM NEAR FORBES Bulman, O.M.B. (1928). In 1927-67. A Monograph of the British Dendroid Graptolites. Palaeontographical Society Monographs, i-ixiv, pp. 1- 97. Bulman, O.M.B. (1938). Graptolithina. In Handbuch der Paldozoologie, in O.H. Schindewolf (ed.), 2D, pp. 1-92. Chapman, A.J., Rickards, R.B. and Grayson, R. (1993). 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N.S.W., 130, 2009 A Holocene History of the Vegetation of the Blue Mountains, New South Wales JANE M. CHALSON! AND HELENE A. MARTIN? '46 Kilmarnock St. Engadine, N.S.W. 2233 * School of Biological, Environmental and Earth Sciences, University of New South Wales, Sydney Australia 2052 (h.martin@unsw.edu.au) Chalson, J.M. and Martin, H.A. (2009). A Holocene history of the vegetation of the Blue mountains, New South Wales. Proceedings of the Linnean Society of New South Wales 130, 77-109. The Greater Blue Mountains Area has been inscribed on the World Heritage list for its exceptionally diverse Eucalyptus communities. Hanging swamps in this region, listed as ‘vulnerable ecological communities’, accumulate sediments that contain the palaeoenvironmental record. Seven of these swamps have been studied, revealing a history of the vegetation, climate and fire regimes. Palynological analysis of each swamp reveals a history of the surrounding vegetation. There are similarities and parallel changes between some of the swamps allowing generalities about the climate of the Holocene to be made. In the early Holocene, about eleven to nine thousand years ago (11-9 ka), the vegetation was more wooded and the climate was probably somewhat warmer and wetter. By the mid Holocene about 6-4 ka, trees were less dominant in the vegetation suggesting that the climate was probably drier. By 3-2 ka, wooded vegetation had mostly returned, and after 2 ka, Baeckea, Leptospermum, Kunzea and Melaleuca species increased somewhat, with further increases in European settlement time, possibly reflecting a reduction or thinning of the wooded canopy. Charcoal analysis of the accumulated sediments suggest that there was more fire in the early Holocene when trees increased the biomass. There was less fire through the mid Holocene when the biomass was lower, but it increased with the return to more wooded vegetation in the late Holocene. In particular, the woody shrubs of Baeckea, Leptospermum, Kunzea and Melaleuca increased with an increase in charcoal, probably because these shrubs benefit from a more open canopy, but they also grew on the swamps hence could deposit charcoal directly into the sediments. Charcoal values are particularly high after European settlement. It is possible that the disruption of Aboriginal burning practices allowed the increased growth of woody shrubs and hence a much greater fuel load. Manuscript received 21 May 2008, accepted for publication 17 December 2008. KEY WORDS: Blue Mountains, Climate change, Fire history, Palynology, Vegetation history. INTRODUCTION The Greater Blue Mountains Area was inscribed on the World Heritage List in December 2000. The Blue Mountains are a deeply incised sandstone plateau rising to over 1,300 m at its highest point. This plateau is thought to have enabled the survival of a rich diversity of plant and animal life by providing a refuge from climatic changes during the recent geological history. It is particularly noted for its wide representation of habitats, from wet and dry sclerophyll, mallee heathlands, as well as localised swamps, wetlands and grassland. Ninety one species of eucalypts are found in the Greater Blue Mountains Area and twelve of these are believed to occur only in the Sydney sandstone region (Australian Government, Department of the Environment and Water Resources, 2007a). The area has been described as a natural laboratory for studying the evolution of the eucalypts (Australian Government, Department of the Environment and Water Resources, 2007a). The steep terrain and sharp environmental gradients have allowed for major evolutionary change in some taxa, resulting in exceptional biodiversity, particularly within the eucalypt communities that dominate the place. Importantly, the evolutionary processes underpinning this diversity are believed to be ongoing, resulting in an evolutionary ‘laboratory’ that is exceptional in the world (Australian Government, Department of the Environment and Water Resources, 2007a). HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION Peat formation on sandstone, the substrateofmost distribution and demonstrable threat has meant that of the Blue Mountains, is very unusual. The hanging _ these hanging swamps are now listed as ‘vulnerable swamps of the Blue Mountains are especially notable | ecological communities’ under the NSW Threatened and have lower sediment loads and accumulate Species Conservation Act of 1995 ((Australian organic matter more slowly than valley swamps and Government, Department of the Environment and swamps along watercourses. They are also easily | Water Resources, 2007b; Sullivan, 2007) eroded with any disturbance. The small geographic Seven swamps in an altitudinal sequence in | QLD | + Swamp, this study GN Other study sites N Ww 33° 00’ —- Be i pi Kings Waterhole Stay is QH2__— Urban areas Area Wee es sydney ers Road = IE ‘S0km Scale S cer a ke a 15 km = & eo S» 33° 15’ — x eee, * Newnes Sw. ihe fe Mountain : *%, Lithgow x Gooches Crater 33° 30' : § es Bell Se aaa aelarmss cee, *., ae 6, ce - ag i Burralow Cr.Sw a. Mt. Victoria ( __ Blackheath @ ee Katoomba : Lawson 7... Penrith BS e. ae wt {x Lakes Sw. : % ce Warrimoo Oval Sw. BSS Pre at loot been a 5 “‘Penmith SS Kings Tableland Sw. Glenbrook "77" coxs Jenolan R. n Notts Sw. \ LAS Warragamba Dam 7 T 7 I 150° 00’ 150° 15’ 150° 30’ 150° 45’ Figure 1. Locality map. 78 Proc. Linn. Soc. N.S.W., 130, 2009 J.M. CHALSON AND H.A. MARTIN the Blue Mountains (Fig. 1) were chosen for a palynological study and are described in Chalson and Martin (this volume). A method to identify Eucalyptus pollen to species was developed (Chalson and Martin, 1995) with the aim of revealing the history of the eucalypt communities of the region. At the beginning of the Holocene, 10,000 years ago, the climate was approaching that of today, but there have been changes through the Holocene (Allan and Lindsay, 1998). The history of the Holocene is thus the history of vegetation very like that of today. THE ENVIRONMENT Geology and geomorphology The Blue Mountains consist of a deeply dissected plateau rising from the Cumberland Plain in the east, along the Lapstone Monocline. Elevation is about 30 m in the east to over 1,000 m in the west. The sedimentary rock units are Triassic in age and curve upwards, from east to west, towards the edge of the Sydney Basin. In the east, Wianamatta Shale outcrops along the side of the Lapstone Monocline. West of the Monocline, the underlying Hawkesbury Sandstone Formation outcrops and further west, underlying the Hawkesbury Sandstone, the Grose Sub-Group of the Narrabeen Group outcrops. The Grose Sub-Group is divided into a number of formations and the ones encountered in this study are as follows: The Banks Wall Sandstone Formation, within which is found the Wentworth Falls Claystone Member, and the basal Burra-Moko Head Sandstone Formation, which is the most prominent cliff-forming unit in the Blue Mountains (Bembrick, 1980). The plateau surface is undulating with small creeks forming upland valleys. In areas where Hawkesbury Sandstone is the underlying rock type, the upland valleys progressively increase in gradient as they incise below the plateau surface and develop steeply inclined V-shaped gorges with only minor benching in the valley sides. To the west, where the Banks Wall Sandstone formation is the underlying rock type, the valley sides and floors slope gently and the streams do not incise but flow across a series of swamps and sandy peat deposits. Eventually, the streams cut through a sandstone layer into claystone or shale when a nickpoint (often a waterfall) is formed (Langford-Smith, 1976). The development of the swamps in these two areas varies enormously. The eastern region supports few swamps which are usually associated with large streams that have a central channel and flowing water. In the western region, there are more swamps and they are developed in broad shallow valleys with no Proc. Linn. Soc. N.S.W., 130, 2009 marked central stream but rather experience a general slow flow of water across the whole area (Langford- Smith, 1976). The climate Maximum temperatures in the Blue Mountains relate strongly to altitude. Average January maxima are highest at the lower altitudes, 29 °C at Richmond and lowest at the higher altitudes, 23 °C at Mt. Victoria. Average minimum temperatures generally decrease from east to west. The July minima range from 3.4 °C at Richmond to —0.8 °C at Lithgow (Table 1). Temperatures as low as —3 °C have been recorded from Katoomba (BoM, 2006; Bureau of Meteorology, 1979). Rainfall patterns relate to elevation and distance from the coast. The average annual rainfall increases from 806 mm at Richmond to 1424 mm at Newnes (Table 1). The driest months are usually July to September and the wettest are December to March (BoM, 2006; Bureau of Meteorology, 1979). Winds from the west or northwest dominate all the year, although there are significant easterly and northeasterly winds during the summer months of November to April. Fogs frequently occur on the higher Blue Mountains, with Katoomba and Mt. Victoria recording an average of 55 and 90 fog days per year, respectively (BoM, 2006). Frosts occur on 35 to 40 days of the year, mostly between April and November. Snow falls most frequently in July and August: Katoomba and Mt. Victoria have and average of 3 and 10 snow days per year, respectively (Bureau of Meteorology, 1979). Soils The quartz-rich sandstones in the area are low in most nutrients, and thus soil and alluvium derived from sandstones are low in nutrients. The soils are mainly lithosols and yellow podzolics with small areas of red and lateritic podzolic soils and sandy alluvial soils in the valleys. Most of the soils are moderately acidic, with pH values of 4.5 to 5. In rugged terrain, rock commonly lies near or at the surface. The soil fertility in the valleys may be higher because of the accumulation of organic matter (Chalson, 1991) Vegetation The vegetation is almost entirely dry sclerophyll woodland and open forest, the “Sydney Sandstone Complex’ (Keith and Benson,1988) with localised swamps in the valleys. There are small patches of tall open forest or wet sclerophyll in specially favourable habitats, such as protected gorges. Heathlands are found in the harshest environments. 79 HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION Table 1. Climatic Averages. Stations are arranged according to altitude. Mean max. temp, hottest month, Station and altitude (m) (Jan.) °C 'Richmond, 19-20 + 29.5 *Penrith, 27 = *Springwood ~400 - *Kurrajong Heights, ~550 - *Lawson, 715 = *Wentworth Falls, ~900 Lithgow (Birdwood St.), 950 DNS Katoomba, 1030 23.1 Mt Victoria, 1064 23.0 *Blackheath PO, 1065 - Lithgow (Newnes Forest 332 Centre), 1050 f Mean Mean min. temp, annual coldest month, x rainfall, (June or July) °C mm 3.4 806 - 786 - 1076 - 1253 : 1260 1409 0.7 860 OES 1398 ley 1061 = 1145 -0.8 1072 1 From BoM (2007). 2 From Bureau of Meteorology (1979) 3 Average of Richmond RAAF and Richmond UWS Hawkesbury Open forest with Angophora costata, Eucalyptus piperita, E. agglomerata and Syncarpia glomulifera dominant is found in sheltered gullies with moist, well-drained soils on the Hawkesbury and Narrabeen Group sandstones. The understorey includes small trees of Allocasuarina torulosa and Acacia elata, with shubs of Hakea dactyloides, Pultenaea flexilis and Dodonaea triquetra. Tall open forest is restricted to the more sheltered gorges and is dominated by E. deanei with Syncarpia glomulifera, Acacia elata, Ceratopetalum apetalum, Callicoma serratifolia and Angophora floribunda. There is a distinctive riparian scrub of Tristaniopsis laurina and Backhousia myrtifolia along the larger water courses (Keith and Benson,1988), Woodland and low woodland with Corymbia gummifera, Eucalyptus sclerophylla and E. oblongata dominant is widespread on ridges and open slopes on shallow, well-drained soils of the Hawkesbury and Narrrabeen Group sandstones. E. punctata, E. piperita and Angophora costata may be present in the more sheltered sites. E. sclerophylla is particularly common on damper soils. The understorey is rich in shrubs of the Proteaceae, Myrtaceae and Fabaceae (Keith and Benson, 1988). There are other woodlands: the “Tablelands Grassy Woodland Complex’ with Eucalyptus dives, E. mannifera, E. eugenioides, E. pauciflora, E. rubida, E. aggregata and E. stellulata the common species. 80 The ‘Snow Gum Woodland’ has FE. pauciflora, E. dalrympleana, E. rubida and E. stellulata dominant (Keith and Benson, 1988). Open heath communities have Eucalyptus stricta, Allocasuarina nana and Leptospermum trinervium, Phyllota squarrosa, Eriostemon obovalis, Epacris reclinata, Dracophyllum secundatum and Gleichenia rupestris dominant. Phyllota squarrosa and Eriostemon obovalis are common in montane heaths whereas Phyllota phylicoides and Eriostemon hispidula are common on the Lower Blue Mountains heath. Many other smaller shrubs are found in these heath communities (Keith and Benson,1988). Closed heath or ‘Newnes Shrub Swamps’ have Leptospermum lanigerum, Baeckea linifolia, Grevillea acanthifolia and Xyris ustulata dominant. They are found in shallow valleys above 1,000 m elevation in swamps, with poorly drained, acid and sandy peat soils. There is a ground cover of sedges including Baloskion australe, Empodisma minus, Lepyrodia_ scariosa, L. anathria, Lepidosperma limicola and small shrubs (Keith and Benson,1988). Closed sedgeland, the ‘Blue Mountains Sedge Swamps’, have Gymnoschoenus sphaerocephalus, Lepidosperma limicola, Xyris ustulata and Baeckea linifolia dominant. These sedge swamps are found at lower altitudes than the closed heath swamps and occupy steep-sided basins (the ‘hanging swamps’). They are intermittently waterlogged and have shallow Proc. Linn. Soc. N.S.W., 130, 2009 J.M. CHALSON AND H.A. MARTIN sandy soils. Many sclerophyllous shrubs form an open heath (Keith and Benson,1988). For a full description of the specific vegetation found at each site, see Chalson and Martin (this volume). Human Occupation The Blue Mountains, especially the lower part, was highly favourable to the hunter-gatherer, (Stockton,1993a). Movement was relatively easy on the ridges, water was not scarce while flora and fauna suitable for food were both plentiful and varied. The rivers were also a source of rock types used for tool making. Campsites with an abundance of worked stone were particularly common in the Lower Blue Mountains. In the Upper Mountains, there were fewer campsites than in the Lower Mountains, but their concentration of flaked stone showed that they have been equally well used. The Central Mountains reveal many rockshelter sites where there were fewer stone artifacts than the Upper and Lower Mountains. However, there was a high concentration of rock art, engravings, paintings and axe grinding grooves. This suggests that the Upper and Lower Mountains were used for survival but the Central Mountains were more of religious and ritual significance (Stockton, 1993a). It is generally presumed that the climate in the Blue Mountains was too severe for year-round occupation during the ice age. However, protected sites such as the rock shelters would have been livable, especially if protected from the bitter westerly winds. (Stockton, 1993b). The oldest signs of occupation in the Blue Mountains were found at Kings Tableland, Wentworth Falls with the oldest date of 22,240 years BP. Walls Cave at Blackheath and Lyre Bird Dell, Leura both yielded dates of more than 12,000 years BP. There were other sites, e.g. Hazelbrook, to 7,200 years BP, Springwood Creek Rock Shelter, from 8,500 years BP up to European times and open sites, e.g. Jamison Creek. Evidence from the Nepean River, at the foot of the Blue Mountains suggests human occupation could go back to 40,000 years BP. In all, there were over 700 Aboriginal sites in the Blue Mountains (Stockton, 1993b; Attenbrow, 2002). With the coming of Europeans, both Europeans and Aborigines avoided each other and early travelers in the Mountains rarely saw any Aborigines. Settlers followed the first crossing of the Mountains in 1813 by Blaxland, Lawson and Wentworth (Breckell, 1993) After some skirmishes about the land the settlers had taken, Aborigines and Europeans co-existed, though not without racist incidents (Smith, 1993). Proc. Linn. Soc. N.S.W., 130, 2009 METHODS Seven swamps in an altitudinal sequence were chosen for study and they are described in Chalson and Martin (this volume). A study of the pollen in surface samples from swamps (Chalson and Martin this volume) provides insights that assist in the interpretation of the pollen spectra from the sediments. The description of the vegetation at each site is also presented in Chalson and Martin (this volume). The swamps were systematically probed to identify the area where accumulating sediments were the deepest, using a Russian D-corer (Birks and Birks, 1980). The sediments and stratigraphy were described using the terminology of Birks and Birks (1980) Samples for radiocarbon dating were taken from a pit where possible, otherwise with repeated use of the D-corer until sufficient sediment was acquired. The standard radiocarbon dates were calibrated using the CalPal (Version March 2007) program. Samples of sediment were taken from the core every 10 cm, or where it was thought there could be a critical change, every 5 cm. For pollen preparations, the core sediments were spiked with Alnus of a known concentration, treated with hydrochloric and hydrofluoric acids to remove siliceous material (Birks and Birks, 1980), oxidised with Schultz solution (a saturated solution of potassium perchlorate in nitric acid), cleared in 10% potassium carbonate and the residue was mounted in glycerine jelly (Brown, 1960). Reference pollen was treated with standard acetolysis (Moore et al., 1991) and also mounted in glycerine jelly. Pollen was identified by comparing grains from the core with a collection of reference pollen. Special attention was paid to pollen of the family Myrtaceae which may be identified to species following the method in Chalson and Martin (1995). Pollen was counted along transects across the slides and tests showed that a count of more than 140 grains adequately sampled the residues. The counts were presented as percentages of the total count and pollen concentrations were calculated for the most abundant pollen groups. Percentages are relative and a change in a single pollen group will affect percentages of all the other groups, but presenting both percentages and concentrations will reveal fluctuations in individual pollen groups. The abundance of charcoal retained on a 150 um sieve, as part of the palynological preparation, was estimated subjectively on a scale of 0 to 8. Counts of microscopic charcoal for a swamp at Kings Tableland showed that the two methods gave similar results, although the microscopic charcoal was more variable (Chalson, 1991). 81 HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION RESULTS Burralow Creek Swamp Burralow Creek Swamp, at 33° 32’S, 150° 36’ 38”E and 310-330 m altitude, is situated in a narrow V-shaped valley and follows the course of the creek for some 3.5 km. The substrate is Hawkesbury Sandstone, but Wiananatta Shale outcrops on the surrounding ridge-tops. The upper reaches of Burralow Creek drain urban areas and farmland areas. An isolated farm adjacent to the swamp was incorporated into the Blue Mountains National Park. Weed growth from this farm is confined to a small area and has not spread into the adjacent bushland. Stratigraphy: Sediments were recovered to a depth of 310 cm. Clayey peat was found down to 10 cm, humic clay at 15-50 cm and humic sandy clay at 60-70 cm. Sand was encountered at 80-260 cm and clay/sand at 260-310 cm. The radiocarbon dates are presented in Table 2. Swamp vegetation and surface pollen: Species of Kunzea and Leptospermum were dominant on the swamp but Restionaceae, Cyperaceae and Selaginella Species were also present (Chalson and Martin, this volume). Surface sample pollen from the swamp (Chalson and Martin, this volume) showed appreciable Leptospermum/Baeckea and a considerable amount of Restionaceae or Cyperaceae in some samples. The fern spore content was low. The pollen record: The pollen spectra from the sediments is presented in Figs 2A, 2B and has been divided into the following zones: 310 to140 cm, no pollen recovered. Zone E, 130 cm, age ? > 1,200 cal yr BP (see Fig. 3 for estimated ages). Angophora floribunda, Eucalyptus spp. and possibly Casuarinaceae pollen were the most abundant of the possible arboreal groups. There was a moderate representation of Poaceae and Selaginella (Fig 2A) and other shrubs and herbs were present in low frequencies (Fig 2B). 120-110 cm, no pollen recovered. Zone D, 100-90 cm, age c. 1,200 —1,000 cal yr BP. . This zone had a very high proportion of Selaginella spores and low proportions of everything else, including tree pollen. The pollen concentrations showed a similar pattern to that of the percentages which revealed a change in the whole pollen spectrum, not only reflecting the addition of a large number of Sellaginella spores to spectra otherwise like that in zone E. Zone C, 80-60 cm, age c. 1,000-800 cal yr BP The Sellaginella content had decreased considerably when compared with the zone brlow, There was a high proportion of Casuarinaceae and Myrtaceae, including Eucalyptus species and the Poaceae content was low. Zone B, 50-20 cm, age c.8700-250 cal yr BP. The Casuarinaceae content had increased and was the highest for the profile. Eucalyptus species and Angophora floribunda were well represented and Leptospemum juniperinum was present in low frequencies. There was a moderate content of Poaceae and Cyperaceae, with a diversity of fern spores. Sellaginella content was minimal. Zone A, 15-0 cm, age c. 250-present, cal yr BP. European Pinus was found in this Zone and there was a high content of L. juniperinum. There was some change in the Eucalyptus species, Casuarinaceae declined. and the Poaceae content was moderate, when compared with the zone below. Charcoal content was low to moderate through most of the profile, with a somewhat higher content at the base of Zone A, the zone of European influence. Table 2. Radiocarbon ages for Burralow Creek Swamp Depth (cm) Material dated Laboratory no. 15-20 Humic clay SUA-2607 50-60 Humic sandy clay SUA-2608 80-90 Sand SUA-2609 95-105 Sand SUA-2610 125-135 Sand SUA-2611 82 Radiocarbon years _—_ Calibrated age (cal yr (yr BP) BP.) 250 +50 3484130 830+60 848+70 1,070450 1068+50 820+50 818450 660455 688450 Proc. Linn. Soc. N.S.W., 130, 2009 J.M. CHALSON AND H.A. MARTIN ‘xipusddy aas ‘uoneja80 oy} ul od4} uatfod ay} Jo vd.41n0s a[quqoid 104 ‘vajoods uatjod dureMg Yae1D MOVING “GZ “WZ SoINSI as soysuiesB ,OL X OF pajunos saiods pue %OZ FS Be feos te Ril feasrvaa] = id t J — : ta es "WS Z | pues ie al Wad [~~ se uajjod jeyo{ = wns ualjod je}oL ee - pe OSF889 | | ey ik it = | : | | | ca a eae TRE area st ieee > a= aaa { | | OZL ? 1 ua}iod oN | 001 = al - 08 | 09 (a ees ov besapn cl Pike : = Teves se ry rr Tan] 77) eesuensnenl =] | fad, ~ » O @ : § £ ££ Pf ff fee 2 g < S > & § § § & €é& Cen eee Si eee g 25h 2 y ga ae sy oF “so o1dosso9e | 2 ¥ g 4 é a S. > § iS a $ é & € G< & oe f S Sos see y & # i f se SS 8 . ri) : 7 Ss S é s $ y Ve ® a 4% yy" S ¥ 83 Proc. Linn. Soc. N.S.W., 130, 2009 84 ponunuos Z ainsi HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION Depth (cm) OOL lo) Some, tic) i= a =) a 28 nig 2 8 a c i= o oO = = ph ™ a = Q ° o 4 P= @ ago 3 2 = ig) er oO a = oO C2) et io} = ° 3 | oo ne) A = 5 = 3 oO & ® te 9) . 2 = roy = rs a = a rom 5 a @ Proc. Linn. Soc. N.S.W., 130, 2009 J.M. CHALSON AND H.A. MARTIN History of the vegetation: Initially, more than 1,200 cal yr BP, there was a mixed tree cover of Myrtaceous species and possibly Casuarinaceae with a moderate Poaceae understorey. Selaginella, was prominent on the swamp. A period of possibly a reduced tree cover followed, with an expanded swamp area with abundant Selaginella about 1.2-1.0 cal ka. Alternatively, if the swamp area was larger, the trees may have been further away, hence they contributed less pollen to the spectrum. The tree cover increased and Selaginella was much reduced by about 1-0.8 cal ka. At this time, the clay content of the sediments increased, perhaps indicating a less energetic water flow. Casuarinaceae became prominent about 0.8-0.25 cal ka with less Myrtaceae, although a diversity of species was identified. Simultaneously, Se/laginella decreased while Cyperaceae and Poaceae increased. In the European zone, there was some change in Eucalyptus species and a big decline in Casuarinaceae while Leptospermum juniperinum became prominent. Fire was a constant factor in the environment, especially in the early part of the European zone. Warrimoo Oval Swamp Warrimoo Oval Swamp, at 33° 43’ 21.44’S, 150° 36’ 58.35”E and 190-200 m altitude, is situated in a V-shaped valley with a stream flowing through it. The substrate is Hawkesbury Sandstone, but Wiananatta shale outcrops on the surrounding ridge-tops. Substantial urban areas occur within a kilometre from the swamp and weed invasion is considerable. Stratigraphy: Total depth recovered was 250 cm. The top 20 cm was peat, then sandy peat down to 50 cm. A layer of sand was found between 50 and 90 cm, then sandy silt down to 200 cm, then sand down to 250 cm when coring stopped (Fig. 4A). The radiocarbon dates are given in Table 3. Swamp vegetation and surface pollen: Species of Baeckea, Kunzea and Leptospermum were dominant on the swamp. Cyperaceae, Juncaceae and Gleichenia species were also present (Chalson and Martin, this volume). The pollen spectra from the surface samples (Chalson and Martin, this volume) contained appreciable Melaleuca, Baeckea/Leptospermum and Gleichenia species. The pollen record: The pollen spectra from the sediments are shown in Figs 4A, 4B. Zone B, 250-130 cm, c. 4,700-2,200 cal yr BP (for estimated ages, see Fig. 5). Abundant Gleichenia denoted this zone, The Myrtaceae content was low, with some of the pollen identifiable to genus/species. There was a consistent content of Casuarinaceae and Haloragis, and Poaceae was almost entirely absent. Zone A, 120 cm to surface, c. 2,200-present cal yr BP. There was very little Gleichenia,, together with an increase in the Myrtaceae and Casuarinaceae content, when compared with the zone below. The Poaceae, Cyperaceae and Restionaceae content was higher and the pollen flora considerably more diverse when compared the preceding zone. Pinus was found down to a depth of 20 cm, thus denoting the European influence, where Baeckea/Leptospermum species increased and Casuarinaceae decreased. The charcoal content was consistently very low in zone B (4.7-2.2 cal ka) and higher in zone A (2.2 cal ka to present). History of the vegetation: From about 4.7-2.2 cal ka, myrtaceous species and Casuarinaceae dominated open vegetation communities. The swamp supported abundant Gleichenia. About 2.2 cal ka, the tree cover of the dryland vegetation increased, with Eucalyptus spp and Leptospermum spp. becoming more diverse and abundant. Casuarinaceae was also more abundant. Gleichenia declined dramatically, but this change was not accompanied by any visible change in the sediments. Fire appears to have been a rare feature of the environment when Gleichenia was dominant. With the change to a more diverse flora and increase of Leptospemum in the swamp community after 2.2 cal ka, fire was more common, particularly in the Table 3. Radiocarbon ages for Warrimoo Oval Swamp Depth (cm) Material dated Laboratory no. 15-25 Peat SUA-2603 120-130 Sandy silt SUA-2604 160-170 Sandy silt SUA-2605 240-250 Sand SUA-2606 Proc. Linn. Soc. N.S.W., 130, 2009 Radiocarbon years Calibrated age (cal (yr BP) yt BP) 730480 738480 2,190+80 2,248+100 2,880+70 3,088+110 4,060+80 4,668+140 85 HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION Se ee ee SPS Seis ° (=) 3 o oS ro) ro} S oO =) o o ' ' b Y n ' 1 oo a mt i ar peice > Peps My q 4 a eRe TET aa ee | H : = "s rs i I S ALL an 5 = : = beers eet | Ses, eee = Con Crap Oo Ye) ~~ ae 1 th = T ieee "Or 35 Be pape 7p See 8] 8 - | oe TPT S hes 9 Bees : of Ly, = “yo, Peo =] 17 o Y i} are iF 1 | i] A] if) i 1] no = Pap * + mop do \} ~ Nagy, eg Yay, Oey Cas, a 4 r {1 5) OS¢., "Pree ‘2 4 1 2 | 1 \ vee bee a oy 7 . So LVS g bro Phy ety ana prong gts Mame MNase 6 2 % 3 area er aes re =) oe \\- — icc ' a sty aT ay a8 2 hp a pr a yg ty 59 1 ET TTT SE ra nag Rig ii i Ming” i Poy : 3 ‘ sNcsonthtaseckiny cles Maar 7 pa) sa a OW Mhoatiet faa naa igg tical Ae | ie jms Cae, 0 : Sip, it , “ag Si = z + 7 team Ae? pe - : ala { *n | r oe Sl Kis I a. | i Af TT Po we q 7 ae he Ero il HI “Tt a wil Ch bios: ee meee) a banca ig. ee Von : To, Ls 4 7 it pera Ae ieee SYS aS SER ator: ie a {— *} | T esl Coa} En A i coat ey, ee, : (a) emg pg egypt 7 o) a a Tape Sen, C856 : “a SS herp aa) ee Pore, mie 3 | rel =z oe ( | | | i - —8 L rs) w 8 ao ah trig Sy a ee tt a = a . ® ae 5 pt i Rote’ OvlFg99'F *xipuoddy 99s ‘uo -8}930A oy} Ul od} UaTjod 34} JO 9d.1NOS s[quqoid 10,7 *e.ajoods uayjod durwas [vaAQ OOWLLILAA “Ap ‘VP Soins 86 Proc. Linn. Soc. N.S.W., 130, 2009 J.M. CHALSON AND H.A. MARTIN ® & rd Sts 2 gf e £ ey 2 § ® © > Fs 2 S 7 2. cians SF 3 # se SF&F EEFEEESB FF SS FE TES EL ES se ST te GS Se GS < 0 Fite dy, Fl
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Proc. Linn. Soc. N.S.W., 130, 2009
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
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Depth (cm)
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J.M. CHALSON AND H.A. MARTIN
Table 5. Radiocarbon ages for Ingar Swamp
Depth (cm) Material dated
30-40 Humic clay with roots
120-130 Sandy humic clay
140-150 Sandy clay
Laboratory no. Radiocarbon years (yr BP.)
BETA 20942 105.1+0.8% modern
BETA 20943 6,460+100
BETA 20944 6,2204100
Calibrated age
(cal. yr BP.)
Modern (<43)
7,428+90
7,188+90
unidentifiable. There was moderate Casuarinaceae
and Poaceae.
The pollen record: The pollen spectra from the
sediments is shown in Fig. 8A, 8B and has been
zoned thus:
Zone C, 150-130 cm, c. 7,000 cal yr BP (see Fig. 9
for estimated ages). Abundant Restionaceae marked
this zone. Eucalyptus piperita, other Myrtaceae
and Casuarinaceae were prominent and there was a
moderate content of Poaceae.
Zone B, 120-40 cm, c. 7,000-?2,200 cal yr BP. There
was greater diversity here and more of tree/large
shrub pollen, viz. E. piperita, Angophora, Melaleuca
&
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60
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100
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Figures 8A 8B. Ingar swamp pollen spectra. For probable source of the pollen type in the vegetation,
see Appendix.
Scale
ram font
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Proc. Linn. Soc. N.S.W., 130, 2009
ES Fe E |
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i
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
Restionaceae decreased but Cyperaceae,
Selagiella and Gleichenia increased
slightly. In the European zone, there
was a Slight decline in Casuarinaceae
and an increase in the swamp species
of Restionaceae and Gleichenia. Fire
was relatively rare about 6 cal ka, but
increased through time, to a peak in the
European period.
Kings Tableland Swamp
Kings Tableland Swamp, at 33° 45”
47” S, 150° 22’ 43” E and about 780-
790 m altitude, is located in the floor
of a steeply sloping small valley off
Queen Victoria Creek. The valley floor
steepens abruptly below the swamp and
a waterfall cascades over a small cliff.
The Banks Wall Sandstone Formation
underlies the swamp and the Wentworth
Falls Claystone outcrops near the base
of the swamp. An area of development
is found less than 1 km to the west
where exotic conifers have been planted
in the gardens.
Stratigraphy: The core sampled 220 cm
of sediments which were peat down to
10 cm, then peaty sand at 15-20 cm,
humic sand at 30-40 cm, peaty silt at 50
cm, humic sand at 60-90 cm, clay/sand
at 100-120 cm and sand at 130-220 cm.
Radiocarbon dates are given in Table 6.
140
Figure 8 continued
styphelioides and Casuarinaceae, when compared with
the zone below. There was a little more Poaceae but
less Restionaceae than in the zone below. Selaginella,
and to a lesser extent, Gleichenia, were
moderate in the base of the zone. 0
European zone
Zone A, 40-0 cm, c. 2,200-0 cal yr BP to
modem. Pinus was found down to 20 cm,
marking European settlement. The dryland 50
flora was similar to the zone below, but tree
species declined with European influence.
Restionaceae and Gleichenia were more
abundant than in the zone below. 400
There was very little charcoal in the
basal zone C, increasing in zone B and
reaching a maximum in the European zone
A.
Zone A
SEEaalaeeearasad Ke ; ;
Zone C ‘ ;
0 “waz 4 6
History of the vegetation: Before 7 cal ka , Age (k yr) ] Radiocarbon date
the vegetation was relatively open, but after @ Calibrated date
about 6 cal ka, the tree cover increased,
especially Casuarinaceae. On the swamp, Figure 9. Ingar Swamp summary diagram
92 Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Table 6. Radiocarbon ages for Kings Tablelands Swamp
Radiocarbon years (yr Calibrated age
BP.) (cal. yr BP.)
a
1.045+0.008 x Modern (< 33)
modern
1,210+70 1,208+90
2,410+70 2,578+130
9,040+80 10,208+130
Depth (cm) Material dated Laboratory no.
15-20 Peaty sand SUA 2656
50-60 Humic sand SUA 2657
80-90 Humic sand SUA 2658
155-160 Fine sand SUA 2659
The swamp vegetation and surface _ pollen:
Leptospermum species were dominant, but Gleichenia
and sclerophyllous shrubs were also found on the
swamp (Chalson and Martin, this volume). In the
surface samples, the Myrtaceae content was low but
Casuarinaceae was well represented (Chalson and
Martin, this volume). The swamp taxa Restionaceae,
Selaginella and Gleichenia were also well represented
and the introduced Pinus was abundant.
The pollen record: The pollen spectra from the
sediments (Figs 10A, 10B) have been zoned thus:
Zone C, 200-90 cm, c. ?>12,000-3,800 cal yr B P (see
Fig. 11 for estimated ages). The Myrtaceae content
was low and Casuarinaceae content moderate (Fig.
10A). Sclerophyllous shrubs and Restionaceae were
well represented (Fig. 10B). Gleichenia and other
fern spores were moderate. Eucalyptus deanei was
found in the basal part of the zone and Banksia in the
upper part.
Zone B, 80-30 cm, c. 3,800 cal yr BP to modern. This
zone had some very high pollen concentrations which
mirrored the spectra of the percentages, suggesting
that the high concentrations result from slow sediment
accumulation rather than the increased input of any
one (or more) particular pollen type(s).
The Myrtaceae pollen proportion remained low
but the Casuarinaceae representation had increased,
when compared with the zone below. The proportion
of Restionaceae and Gleichenia had decreased,
but Cyperaceae and Selagiella had increased, in
comparison with the zone below. Sclerophyllous
shrubs were also well represented in this zone.
Zone A, 0-25 cm, modern. Pinus was found here,
delimiting the European zone. The myrtaceous
content had increased a little, especially Melaleuca.
Casuarinaceae and Restionaceae decreased somewhat
but Gleichenia increased considerably, when
compared with the zone below.
The charcoal content was low to moderate in zones C
Proc. Linn. Soc. N.S.W., 130, 2009
and B, and increasing in the modern zone A.
History of the vegetation: The dearth of myrtaceous
taxa, predominance of Casuarinaceae and the diversity
and relative abundance of the shrubby taxa suggests a
heathland, given that the two species of Casuarinaceae
found in the region today, A//ocasuarina distyla and
A. nana, are shrubs/small trees. The swamp flora
was dominated by Restionaceae throughout, with
Gleichenia becoming prominent in modern times.
Myrtaceae remains low until modern times, suggesting
the surrounding vegetation remained relatively open.
The charcoal content was relatively low until modern
times, suggesting less fire activity, or lesser fuel to
burn, until European times.
Katoomba Swamp
Katoomba Swamp, at 33° 43’ 03” S, 150° 19’ 18”
E and 950 m altitude, is located in a small, shallow
valley which is a tributary of Gordon Creek (Chalson
and Martin, this volume). The Banks Wall Sandstone
Formation underlies the swamp and the Wentworth
Claystone Member outcrops near the base of the
swamp, probably impeding drainage. -
This swamp is surrounded by urban development.
There is evidence of drainage ditches and a sealed road
runs across the swamp. Much of it is (or has been)
used for yards for light industry and horse paddocks.
Housing extends to the edge of the swamp.
Stratigraphy: Two cores were necessary to recover
sediments spanning the whole of the Holocene. Core
1 consisted of (1) dark greyish brown or dark brown
silty clay/humic clay/clay with roots, 0-20 cm, then
(2) dark greyish brown, black, or dark grey silty or
sandy clay at 25-80 cm, followed by (3) dark grey
sand at 85 cm, (4) dark grey clay at 90 cm, (5) dark
greyish brown or dark grey sandy or silty clay at 95-
115 cm, (6) dark grey sand at 120 cm and (7) dark
grey sandy clay at 125-130 cm.
The stratigraphy of core 2 consisted of (1) dark
greyish brown, dark grey or dark brown silty clay at
0-30 cm, then (2) dark grey or dark brown sandy clay,
93
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
charcoal
ge
1 4 7 a ia)
yee = =
a ~~
rs a
H
id ~
= ies,
wu Oe
we siay gdp s Ofna aa ae aiewip cop wan xgowent att Skasa oh ie ese nee oan
155 x10° 163x107}
F B
80 . > THe on tne ge Pn 2 Serer Sree see < scores ie C
270 xi0* | 2,578+130 -
100 lee oe C Eee a
120 = = : : ie Wee . i
140 ee 3 - ; —
160 i - . = : 10,2084130.
180 _ : —— e — Le is a.
7 E
| Sand [7] Sit [-=_] Clay boats
Figures 10A, 10B. Kings Tableland Swamp pollen spectra. For probable source of the pollen type in the vegetation, see Appendix.
Proc. Linn. Soc. N.S.W., 130, 2009
94
J.M. CHALSON AND H.A. MARTIN
SED
Figure 10 continued
35-40 cm, followed by (3) dark greyish brown sand at
42-48 cm and (4) dark grey clay or sandy clay at 50-
55 cm. Radiocarbon dates are presented in Table 7.
The swamp vegetation and surface pollen: The
moss Dawsonia, and species of Cyperaceae and
Juncaceae were dominant on the swamp. Kunzea
and Leptospermum species were also dominant and
many sclerophyllous shrubs were found on the edge
of the swamp, but the natural vegetation was highly
disturbed here (Chalson and Martin, this volume).
Poaceae (both native and introduced species) was
the dominant pollen type in the surface samples,
reflecting the urbanisation and the disturbance at the
site. Pinus pollen was also present in appreciable
amounts. Total Myrtaceae pollen was moderate
and Casuarinaceae pollen was low. The swamp
taxa, Restionaceae, Cyperaceae, Selaginella and
Proc. Linn. Soc. N.S.W., 130, 2009
Gleichenia were present in low proportions (Chalson
and Martin, this volume).
The pollen record: Pollen recovery from the cores
was good and some very high concentrations were
found, especially in the clay (Figs 12A, 12B). The
cores were zoned thus:
Core 2, Zone D, 55-0 cm, c. 12-11,000 cal yr BP (see
Fig. 13 for estimated ages). The Myrtaceae content
was low but Eucalyptus oreades and E. pauciflora
had been identified. Casuarinaceae and Poaceae
representation was moderate and Restionaceae
was high (Fig. 12A). Asteraceae/Tubuliflorae and
Ericaceae were prominent amongst the herbs and
shrubs (Fig. 12B). The charcoal content was moderate
throughout.
95
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
1 nm Nay
ie |
Ny 1] Zone A |4 ‘a.
| £ ios
it * at
= 255
5 ZoneB i:
t= y n
| ,
a |
oO | 4 1
1004 OF
| 25
| Zone C
i *
150+ ae
? re
i ”
6
Age (k yr)
i Radiocarbon date
@ Calibrated date
Figure 11. Kings Tableland Swamp summary diagram.
Core 1, zone C, 130-75 cm, c. 6,200-4,000 cal_yr
BP (for estimated ages, see Fig 13). The Myrtaceae
representation was very low, lower than in the zone
below, and Eucalyptus species were not recorded from
most samples. Casuarinaceae representation was low
also, Poaceae was moderate and Restionaceae high,
all fairly similar to the zone below.
Core 1, zone B, 70-30 cm, c. 3,100-?1,500 cal yr BP.
The Myrtaceae content had increased and Eucalyptus
oreades was present through the zone, and this was
the most notable difference when compared with the
zone below. Casuarinaceae abundance was moderate
and the Poaceae representation had decreased when
compared to the zone below. Restionaceae abundance
was a little less than in the zone below, decreasing
further towards the top of the zone. Haloragis and
Grevillea acanthifolia were prominent amongst the
herbs and shrubs.
Coren zonesAce25-0ucmcan eo 00Rcalmyme Pato
present. Pinus was consistently present, denoting the
European zone. Total Myrtaceae and Casuarinaceae
Table 7. Radiocarbon ages for Katoomba Swamp
Depth (cm) Material dated Laboratory no.
Core 1,125-130 Sandy clay Beta 24545
Core 2, 0-5 Silty Clay Beta 24547
Core 2, 50-55 Sandy Clay Beta 24546
96
representation were low, decreasing somewhat from
the base, but E. oreades and A. floribunda were
found throughout the zone. Poaceae pollen increased
markedly from the base of the zone but Restionaceae
was very low at the very base, then virtually absent
from the rest of the zone. Cyperaceae increased a little
and Asteraceae/Liguliflorae was present throughout
the zone.
The charcoal content was very low in zone C, then
low through the rest of the core, with an occasional
moderate value.
History of the vegetation: There was an open or
sparse tree cover about 11-12 cal ka. By 6-5 cal ka,
the site appears to have been almost treeless. About 4
cal ka, E. oreades returned to the site which became
wooded once again. Restionaceae was dominant on
the swamp and Poaceae was moderately common
until 3 kyr BP, after which, both declined. In the
European zone, Poaceae increased dramatically,
no doubt reflecting urbanisation. At the same time
Restionaceae decreased and almost vanished from
the swamp. E. oreades remained dominant but it
Radiocarbon years Calibrated age
(yr BP) (cal. yr BP)
5,450+80 6,288+100
11,0304£130 12,998+120
10,570£100 12,558+170
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
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Proc. Linn. Soc. N.S.W., 130, 2009
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
Figure 12 continued
decreased, along with Casuarinaceae in the time of
the Europeans. Fire activity was low to moderate
through most of the time.
Newnes Swamp
Newnes Swamp, at 33° 22’ 57” S, 150° 13’ 20” E
and 1,060 m altitude, is located in a shallow hanging
valley with pine plantations in close proximity.
Regular burning maintains fire breaks for the young
pine plantations. The swamp is underlain by the
Burra-Moko Head Sandstone Member of the Banks
Wall Sandstone Formation which has thin claystone
interbeds, and it is likely that one of these clay layers
impedes drainage and hence maintains the swamp.
Swamp stratigraphy: The core sampled 90 cm of
sediment. Clay or peat with roots was found down
98
to 20 cm, then sandy clay down to 35 cm, followed
by sand to 55 cm, then sandy clay with roots down to
65 cm, then silty clay to 75 cm, and finally sand or
sandy clay in layers to 90 cm. Radiocarbon dates are
presented in Table 8.
The swamp vegetation and surface pollen: Banksia and
Kunzea were dominant and Baeckea, Leptospermum,
other sclerophyllous shrubs, Cyperaceae and Poaceae
were also present on the swamp (Chalson and Martin,
this volume). There was appreciable Myrtaceae
pollen in the surface samples, but Restionaceae and
Gleichenia were dominant in the surface pollen
spectra. Pinus was present but not abundant. (Chalson
and Martin, this volume).
The pollen record: Pollen recovery from the core
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
50
Depth (cm)
100
0 2
6
Age (k yr)
J Radiocarbon date
@ Calibrated date
Figure 13. Katoomba Swamp summary diagram.
was good and there was some exceptionally high
concentrations, especially in the clay at 60-70 cm.
The core was zoned thus (Figs 14A, 14B):
Zone D, 90-55 cm, c. 11,000-7,5 00 cal yr BP (see
Fig. 15 for estimated ages). Myrtaceae pollen was
low, but Eucalyptus pauciflora/rubida had been
identified. Casuarinaceae was also low at the base of
the zone, increasing upwards (Fig 2A). Asteraceae/
Tubuliflorae and Chenopodiaceae were prominent
amongst the herbs and shrubs (Fig. 14B). Poaceae
and Restionaceae were well represented.
Zone C, 50-40 cm, c. 7,500-1,800 cal yr BP There
was very little Myrtaceae pollen, with only one
record of a Eucalyptus species. Casuarinaceae pollen
increased, Haloragis was moderate and Poaceae and
Restionaceae were reduced when compared with the
Table 8. Radiocarbon ages for Newnes Swamp
Depth (cm) Material dated Laboratory no.
20-25 Sandy clay SUA 2648
35-40 Sandy clay SUA 2649
50-55 Sand SUA 2650
77-83 Silty clay SUA 2651
87-93 Sand SUA 2652
Proc. Linn. Soc. N.S.W., 130, 2009
preceding zone.
Zone B, 35-25 cm, c.1,800-?1,000 cal yr BP. Melaleuca
representation was significant, Casuarinaceae had
decreased, the shrubs were well represented, and
Poaceae and Restionaceae remained low when
compared with the previous zone.
Zone A, 20-0 cm, ?1,000 cal yr BP to present.
Melaleuca continued to be the most significant
of the Myrtaceae, Styphelia and Haloragis were
appreciable, Poaceae remained low and Restionaceae
was somewhat greater than the zone below. Pinus
was present throughout the zone, denoting European
activity.
The charcoal content was moderate in zone D,
extremely low in zone C, and moderate to high in
zones B and A.
Radiocarbon years Calibrated age (cal. yr
(yr BP) BP)
1,090+70 1,098+80
1,930+£70 1,948+80
6,650+100 7,588+80
9,820+90 11,398+130
9,640+80 11,038+160
99
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
y, ~
“era, ae * 2
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es 2 a rs a >
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00) | c=) Ou.
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= * oe b secret oereore as
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107
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
APPENDIX A
Pollen type name on the pollen diagrams and the probable source in the vegetation.
Name of pollen type. Major
pollen groups (A diagram)
Podocarpus
Pinus
Angophora/Corymbia
Eucalyptus/Melaleuca
Melaleuca styphelioides
Leptospermum/Baeckea
Tristaniopsis
Unidetified Myrtaceae
Casuarinaceae
Poaceae
Restionaceae
Cyperaceae
Selaginella
Gleichenia
Other fern spores
Probable source in the vegetation and ecological inference.
From PlantNet (2007)
Probably Podocarpus spinulosus: sclerophyllous shrub/small tree
Pinus sp(p). Introduced: Pollen input from urban/forestry areas.
Species within the two genera: sclerophyll woodland
Species within the two genera sclerophyll woodland/forest
Melaleuca styphelioides: moist stream bank habitat
Species within the two genera: ?mainly swamp communities
Tristaniopsis spp : moist habitats in sclerophyll communities
All pollen types not identifiable further
Casuarina, Allocasuarina sp(p): A. distyla and A. nana in this study
Native and exotic species in the family: open situations, dryland
and swamp species
All species in the family: swamp and dry land species
All species in the family: swamp and dry land species
All species in the genus: damp sites, edge of swamp
Gleichenia sp(p): damp sites, edge of swamp
Other ferns: many possible species
Names of shrubs and herbs (B diagrams)
Grevillea acanthifolia
G. sphacelata
Grevillea
Hakea
Persoonia pinifolia
Persoonia
Symphionema montanum
Banksia
Other Proteaceae
Acacia
Styphelia
Monotoca
Other Ericaceae
Rutaceae type
Pimelea
Plantago
Haloragis
Other tricolporate grains
Podocarpus
Micrantheum
Myriophyllum
Asteraceae/Liguliflorae
Asteraceae/Tubuliflorae
Chenopodiaceae
Shrub: swampy areas, sand or peat
Shrub: heath, dry sclerophyll forest
Grevillea sp(p): sclerophyllous understorey
Hakea sp(p): sclerophyllous understorey
Shrub: heath, dry sclerophyll forest
Persoonia sp(p): sclerophyllous understorey
Shrub: heath or dry sclerophyll forest, wet or dry situations
Banksia sp(p): sclerophyllous understorey
Other taxa in the family: sclerophyllous understorey
All species in the genus
Styphelia sp(p): sclerophyllous understorey
Monotoca sp(p): sclerophyllous understorey
Other taxa in the family: sclerophyllous understorey
All taxa in the family sclerophyllous understorey
Pimelea sp(p): sclerophyllous understorey
Plantago sp(p): native and introduced herbs
Haloragis/Gonocarpus sp(p): Damp sites, sclerophyllous understorey
Probably shrubs and herbs
Probably Podocarpus spinulosus: sclerophyllous shrub/small tree
Shrub: heath and dry sclerophyll forest, sandy infertile soils
Mainly aquatic herbs, also on damp ground around water bodies
Fenestrate-grained taxa in the subfamily Liguliflorae: herbs
Echinate-grained taxa in the subfam. Tubuliflorae: shrubs and herbs
Ruderals, salt tolerant
108
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
APPENDIX B
Myrtaceae Pollen type name on the pollen diagrams and the probable source in the vegetation.
Name on the pollen
diagrams
Angophora costata
Probable source in the vegetation and ecological inference.
From PlantNet (2007)
Deep sandy soils on sandstone
Angophora floribunda
A. costata x floribunda
Angophora
Baeckea/Leptospermum
Callistemon
Eucalyptus deanei
E. eugenioides
E. fibrosa
E. oblonga
E. oreades
E. pauciflora/E. rubida
E. piperita
E. racemosa
Eucalyptus type B
Eucalyptus type C
Eucalyptus type D
Eucalyptus type K
Eucalyptus type M
Eucalyptus/Melaleuca
Kunzea
Leptospermum
juniperinum
L. polygalifolium
Melaleuca ericifolia
M. styphelioides
Melaleuca type B
Melaleuca type C
Melaleuca
Myrtaceae type C
Myrtaceae type D
Unidentified Myrtaceae
Usually on deep alluvial soils
Some species in swamp/moist habitats, also dryland species
Dry sclerophyll communities, some swamp species
Tall wet forest, sheltered valleys, deep sandy alluvial soils
Dry sclerophyll or grassy forest, on deep soils
Wet or dry sclerophyll forest, on shallower, somewhat infertile soils
Dry sclerophyll woodland, on extremely infertile, sandy soils
Wet or dry scleropnhyll forest, on poor skeletal or sandy soils
Grassy or dry sclerophyll woodland, on cold flats.
Dry sclerophyll forest/woodland, moderately fertile, often alluvial
sandy soils
Dry sclerophyll woodland, on shallow infertile soils
)
)
) For definition of Eucalyptus pollen types, see Chalson (1991)
)
)
Species within the two genera: sclerophyll woodland/forest
Understorey sclerophyll forest, moist depressions
Swamp, heath and sedgeland, on sandy peat soils
Dryland habitats and moist depressions
Heath and dry sclerophyll forest, streambanks and coastal swamps
Moist situations, often stream bank habitats
)
) For definition of pollen type, see Chalson (1991)
For definition of Melaleuca pollen types, see Chalson (1991)
)
) For definition of pollen types, see Chalson (1991)
All myrtaceous pollen types not identifiable further
Proc. Linn. Soc. N.S.W., 130, 2009 109
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Modern Pollen Deposition Under Vegetation of the Blue
Mountains, New South Wales
JANE M. CHALSON! AND HELENE A. MARTIN?
'46 Kilmarnock St. Engadine, N.S.W. 2233
* School of Biological, Environmental and Earth Sciences, University of New South Wales, Sydney Australia
2052 (h.martin@unsw.edu.au)
Chalson, J.M. and Martin H.A. (200x). Modern pollen deposition under vegetation of the Blue
Mountains, New South Wales. Proceedings of the Linnean Society of New South Wales 130, 111-137.
Pollen was extracted from surface samples of swamp sediments and soils under various types of vegetation
in the catchments of these swamps. The pollen assemblages in these surface samples were compared with
the floristic composition of the vegetation to provide a means of interpreting the assemblages of fossil
pollen retrieved from the swamp sediments.
The surface pollen assemblages reflected the local vegetation, indicating more/less tree cover, swamp and/
or adjacent dryland environment and local flora diversity. All the evidence pointed to very local deposition
and little long distance dispersal of pollen. A number of different units may be defined within the one major
vegetation type, dry sclerophyll forest/woodland 1n this case, but the floristics of the units are too similar to
allow discrimination of them from their modern pollen assemblages.
Manuscript received 21 May 2008, accepted for publication 17 December 2008.
KEYWORDS: Blue Mountains, local pollen deposition, long distance pollen dispersal, modern pollen
deposition, pollen spectra.
INTRODUCTION
Pollen is deposited in sediments by the
contemporaneous vegetation, but a number of factors
affect the representation of each taxon in the sediments
so that it is not possible to relate a fossil pollen
assemblage in a deposit directly to the vegetation
that produced it. Pollen productivity, dispersal and
preservation are the main factors that influence
representation of a taxon, and each of these factors
are in turn influenced by the local environmental
conditions. Pollen deposited from under known plant
communities, however, may be used to characterize
that community and hence assist in the interpretation
of pollen spectra recovered from swamp sediments.
The nature of pollen deposition of individual taxa
may also be deduced from the surface pollen spectra.
Sites for a study of the history of the vegetation
were chosen from swamps in an altitudinal sequence
in the Blue Mountains (Fig. 1). These sites are situated
on a relatively uniform substrate, sandstone, within
dry sclerophyll woodland/open forest. Observations
of modern pollen deposition are reported in this
paper, and the Holocene history of the vegetation
from the swamps is reported in Chalson and Martin
(this volume).
THE STUDY SITES
The Blue Mountains are a deeply dissected
plateau rising from the Cumberland Plain in the
east. The plateau surface is undulating and small
creeks form upland valleys. Where the underlying
rock type is Hawkesbury Sandstone, the upland
valleys become incised and develop into V-shaped
gorges. In the west where rock type is Banks Wall
sandstone, the valley sides and floors slope gently
and the streams flow through a series of swamps
(Chalson, 1991).
The swamps chosen for study are as follows
(see Fig. 1) and the species found at each site are
listed in Appendix 1:
Burralow Creek Swamp, at 33° 32’S, 150° 38’E and
310-330 m altitude, is anarrow swamp that follows the
creek for some 3.5 km. The upper end of the swamp
is 2 km southeast of Kurrajong Heights. The core site
is | km downstream from the northern end. There are
few cleared areas near the swamp, the nearest being
over 2 km away.
The vegetation around Burralow Creek is open
forest, woodland and swamps (Keith and Benson,
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
33° 30’
sy Mt. Victoria
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Mediow Bath &
Study area
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33° 45’ ~
pees
Jenolan R.
~
150° 00’ 15°
0 Scale 15 km
© &
Urban area
Katoomba Sw:-* Wood NympHs Dell Warrimoo
Kings
Tablelands Sw.
5 ~~" Notts Sw. ~~, \
-, ae > Ce ae Sw.
Oval Sw.
Murphys Glen a es
Ingar Sw. Glenbrook —
Yoo
Warragamba Dam
WP agieg
30 150° 45)
%e Swamp, this study
ve Other study site
& Other surface sample
Figure 1. Locality map
1988). Angophora bakeri, A. costata, Corymbia
eximia, Eucalyptus eugenioides, E. multicaulis, E.
paucifiora and E. radiata are locally dominant with
a few kilometers of the swamp. The surface of the
swamp supports an open heathland of Leptospermum
polygalifolium, L. trinervium and _ Eleocharis
sphacelata. Nomenclature follows Harden (1992;
1993; 2000; 2002) and PlantNet (2006)
2
Warrimoo Oval Swamp, at 33° 43’ 21.44”S, 150° 36’
58.35”E and 190-200 m altitude, is approximately 1.5
km east of Warrimoo Post office and 0.4 km south
of Warrimoo Oval. There are substantial urban areas
within a kilometer of the swamp and weed invasion
is considerable.
The vegetation is mainly woodland with
some open forest and swamp communities (Keith
and Benson, 1988). Locally, Angophora bakeri,
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Eucalyptus pauciflora and E. radiata are dominant.
The swamp surface supports an open heathland with
Leptospermum spp.
Notts Swamp, at 33° 48’ 35.44” S, 150° 24’ 27.66”
E and about 682 m altitude is approximately 12 km
south-southeast of Wentworth Falls and to the west of
Notts Hill. The lower third of the swamp is used as a
market garden, but there is no sign of disturbance or
weed invasion at the study site. There is no indication
of European activities in the catchment upstream of
the study site and the nearest settlement is some 7 km
to the north-northeast.
The major plant community is open woodland
and there is a little open forest and some swamps
(Keith and Benson 1988). Eucalyptus eugenioides, E.
multicaulis, E. piperita, E. racemosa and E. sieberi
are locally dominant. The swamp supports a closed
sedgeland of Gymnoschoenus sphaerocephalus,
Leptospermum trinervium and Baloskion australe.
Ingar Swamp, at 33° 46’ 11.65” S, 150° 27’ 22.92” E
and 584m altitude, is approximately 8 km southeast
of Lawson. European settlement is some five km to
the northeast, along the highway, and includes some
very large, old conifer trees.
The vegetation is mainly woodland with
Corymbia gummifera, Eucalyptus oblongata, E.
piperita, E. pauciflora, and Angophora costata
dominant locally. Open forest in gorges along
the creeks is dominated by E. eugenioides, E.
sclerophylla, Tristania neriifolia and Angophora
costata. The swamp community is a closed sedgeland
of Gymnoschoenus sphaerocephalus, Leptocarpus
tenax, Baumea sp., Chorizandra sp., Baloskion
australe and, towards the edge, Hakea teretifolia, H.
dactyloides and Leptospermum lanigerum.
Kings Tablelands, at 33° 45’ 47” S, 150° 22’ 43” E and
about 780-790 m altitude, is located in small valley
off Queen Victoria Creek. It is about 0.6 km east of
Queen Victoria Memorial Hospital near Wentworth
Falls. An urban area is found less than | km to the
west where exotic conifers have been planted in the
gardens.
The vegetation is mainly open forest around the
study site, with woodland on the ridges and closed
sedgelands in the swamps (Keith and Benson, 1988).
Locally, Eucalyptus dives, E. oreades, E. sieberi
and E. piperita are dominant in the open forest and
Corymbia gummifera, E. racemosa and E. sieberi are
dominant in the woodland. On the exposed plateau
to the northeast, the dominants in an open heathland
are Allocasuarina distyla, E. ligustrina, E. stricta,
Proc. Linn. Soc. N.S.W., 130, 2009
Banksia serrata and Hakea teretifolia. The dominants
on the swamp are Leptospermum juniperinum and L.
grandiflorum.
Katoomba Swamp, at 33° 43’ 03” S, 150° 19’ 18” E
and 950 m altitude, is 1 km east northeast of Katoomba
Post Office and 1 km west of Leura Post Office. This
swamp is surrounded by urban activity, with drainage
ditches and a sealed road running across the swamp.
Much of the swamp is (or has been) used for yards for
light industry and horse paddocks. Housing extends
to the edge of the swamp.
Most of the area around the swamp has been
cleared but there are a few remnant pockets of
Sandstone Plateau Forest (Keith and Benson, 1988)
remaining. Eucalyptus acmenoides, E. oreades, E.
stellulata, E. oblongata and E. sieberi are dominant.
The understorey is problematic as the remnant stands
are heavily weed infested.
Little remains of the original vegetation over
the swamp surface and species of Poaceae are
predominant. A small patch of swamp edge vegetation
forms a dense thicket of Leptospermum juniperinum
and L. scoparium.
Newnes Swamp, at 33° 22’ 57” S,150° 13’ 20” E and
1,060 m altitude, is within a forestry area with pine
plantations. Regular burning maintains fire breaks.
Woodland communities are found around the
swamp (Benson and Keith, 1990) but the shrub layer
has been much reduced by frequent burning. Shrubs
remaining on the swamp include Leptospermum
trinervium and Grevillea acanthifolia. A ground cover
of grasses is found in all but the wettest areas where
Juncaceae and Restionaceae are dominant.
METHODS
The vegetation units at each site were determined
from maps in Benson (1992), Keith and Benson
(1988) and Benson and Keith (1990). Each site was
visited, the vegetation checked with the maps and as
many species as possible were identified in each of
the vegetation units. Since palynology cannot reveal
the structure of the vegetation, the focus of survey
was on the species list. Dominance was determined
subjectively from the abundance of the species
Samples from the surface of the soil, or where
possible, from moss polsters, were collected from the
centre of the swamp, the swamp edge and the plant
communities adjacent or local to, the swamp sites.
Samples were taken from at least 100 m away from
community boundaries where possible. The sample
types and vegetation are listed in Table 1 and the
113
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Table 1 Surface samples used for pollen spectra presented in Figs 2 and 3. Codes for vegetation
map units are from Keith and Benson (1988).
; Vegetation .
STIRS Vegetation e Sample material
sample no. map unit
Burralow Creek
1 Open sedgeland mid-swamp 28a Soil
2 Open sedgeland mid-swamp 28a 0 cm core
3 Swamp fringe 28a Soil
4 Low Woodland 10ar Soil
5 Open forest 10ag Soil
Warrimoo Oval
6 Closed sedgeland mid-swamp 26a Soil
7 Closed sedgeland mid-swamp 26a 0 cm core
8 Closed sedgeland swamp fringe 26a Soil
9) Low woodland 10ar Soil
Notts
10 Closed sedgeland mid-swamp 26a Soil
11 Closed sedgeland swamp fringe 26a Soil
Ingar
12 Closed sedgeland mid-swamp 26a Soil
IL) Closed sedgeland swamp fringe 26a Soil
14 Low woodland 10ar Soil
15 Low woodland 10ar Soil
Kings Tableland
16 Closed sedgeland mid-swamp 26a 0 cm core
iW, Closed sedgeland swamp fringe 26a Soil
18 Low woodland 10ar Soil
19 Low woodland 10ar Soil
20 Open forest 91 Soil
Dil Open forest 91 Soil
22 Open heath Dalits Soil
Katoomba
73 Closed sedgeland mid-swamp 26a Soil
24 Closed sedgeland swamp fringe 26a Soil
1) Open forest 91 Soil
26 Open forest 91 Soil
Newnes
ZT Closed heath mid-swamp 20a Moss
28 Closed heath swamp fringe 20a Moss
29 Woodland 10f/lla Moss
30 Woodland 10f/lla Moss
31 Woodland 10f/lla Soil
32 Woodland 10f/lla Soil
33 Open heath 21d Soil
34 Open heath ZN Soil
35 Forest 10f Soil
36 Forest 10f Soil
Murphys Glen
37 Tall open forest 6c Soil
38 Tall open forest 6c Soil
Wolgan
39 Open woodland lla Soil
40 Open woodland lla Soil
Wood Nymphs Dell
4] Open forest 10ag Soil
Medlow Bath
42 Open forest 91 Soil
114 Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
study sites are shown in Fig. 1
Six to ten sub-samples were taken from each
plant community over a transect of approximately 20
m. The sub-samples were mixed together to reduce
the possible over-representation of any one species
due to close proximity to an individual plant (Chalson,
1991).
The samples were treated with hydrochloric and
hydrofluoric acids to remove siliceous material (Birks
and Birks, 1980), oxidised with Schultz solution (a
saturated solution of potassium perchlorate in nitric
acid), cleared in 10% potassium carbonate and the
residue was mounted in glycerine jelly (Brown,
1960).
Pollen was identified by comparing the grains
with reference pollen treated with standard acetolysis
(Moore et al., 1991). Grains were counted along
transects across the slides and tests showed that a
count of 140 grains adequately sampled the residues.
The counts of each pollen type were presented
as percentages of the total count on the pollen
diagrams.
RESULTS
Fig. 2 presents the pollen spectra from vegetation
on the swamp surface and at the edge of the swamp,
and Fig 3. presents spectra from the dry-land
communities in the surrounding vegetation. Table 2
presents the name on the pollen diagram, the probable
source of the pollen in the vegetation and ecological
inference.
Preservation, although adequate, was not
good enough for the identification of Eucalyptus
species beyond broad groups (Chalson and Martin,
1995). The pollen from moss polsters may be better
preserved than that from the soil, but moss polsters
were not common and usually dried out severely
in the forest environment, hence soil samples were
usually collected in all but the dampest areas.
Exotic Pinus is present in all samples (Figs 2A,
3B) and values are highest at sites near urban areas
(Kings Tableland, Katoomba). Surprisingly, Pinus
values are not high at Newnes, in the forestry area
with pine plantations, but the pines were very young
at the time of this study.
Angophora/Corymbia and Eucalyptus/Melaleuca
have been identified in low frequencies in some of
the samples which were better preserved. Melaleuca
styphelioides has been identified in some of the
swamp samples (Fig. 2A) where counts may be high.
M. styphelioides was not found during the survey of
the vegetation, but it may be grown in gardens. The
Proc. Linn. Soc. N.S.W., 130, 2009
highest count at Warimoo Oval Swamp is close to
substantial urban areas. Leptospermum/Baeckea has
been identified from some swamp samples (Fig. 2A)
where counts may be considerable. Leptospermum
spp. are often dominant in the swamp communities
(see Appendix 1)
The unidentified Myrtaceae group is larger than
the other groups of Mytaceae and counts from the
swamp samples are the lowest of all. The woodland
or forest samples from the borders of the swamp (Fig.
2A) all have higher counts than the swamp samples.
Frequencies in samples from the dry-land vegetation
(3A) are much higher than those from swamps. Lack
of specific identification was generally due to poor
preservation.
Casuarinaceae frequencies are usually low, with
a few higher values. The highest value (Fig. 3A)
comes from heathland vegetation.
Poaceae frequencies are generally low and the
high values are associated with urbanisation and
disturbance (Katoomba, Fig. 2A).
Restionaceae frequencies are variable but most
of the high values are found in the swamp samples.
Cyperaceae has not been recorded from many samples,
and where it is present, frequencies are generally low,
with the few higher frequencies being found in the
swamp samples.
Selaginella is present in a few samples and
appreciable frequencies may be recorded in some
swamp samples. Gleichenia may be present in
appreciable frequencies in some swamp samples
also. Other fern spores are usually recorded in low
frequencies and are more common in the dry-land
samples.
Table 2 also lists the likely environmental
indication of the pollen groups on the diagrams,
but this is difficult, given that a group may include
many possible species. For example, the families
Restionaceae and Cyperaceae include both swamp
and dry-land species, but the species in the vegetation
and patterns of high pollen frequencies on the
diagrams may indicate the nature of the environment
when considered together. Thus the species of
Restionaceae and Cyperaceae found in the local
vegetation (Appendix 1) are almost entirely species
of swamps or damp places (Table 2).
DISCUSSION
There are many indications that the pollen
recovered from the surface samples was
produced mainly by the local vegetation and
thus the pollen spectra can indicate the type of
115
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
iy
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a5 Ro tty
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3 Swamp edge
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11 Swamp edge
12 Mid swamp z |
13 Swamp edge
18 Woodland
16 Mid swamp iz L
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Figure 2A. The pollen spectra from plant communities associated with swamps within major pollen groups. The Sample
number (extreme left hand side) refers to the sample in Table 1.
Proc. Linn. Soc. N.S.W., 130, 2009
116
J.M. CHALSON AND H.A. MARTIN
& &
Scale . 20% of total count 2 Ss = e
QD 5
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Burralow Creek Swamp ab
7 Mid swamp | | | i L j } ! '
2 Mid swamp | | is \ | r
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4 Woodland |= at ed |
oodian Pee Se dL) 4 b
Warrimoo Oval Swamp
6 Mid swamp | + | L iL |
7 Mid swamp = i | | |
8 Swamp edge H aeety |
9 Woodland | - | L L lj r [ |
Notts Swamp
10 Mid swamp | L | | | { ' |
41 Swamp edge | | | | i [ f | | | [ |
Ingar Swamp
12 Midswamp | ] | | |
13 Swamp edge | | | | L { L L | r i
; ees
Kings Tableland Swamp
18 Woodland ‘ | i i
16 Mid swamp | r | - | | | [
17 Swamp edge ig | ri - oP Perks
21 Fores | ee Re de bai
Katoomba Swamp
23 Mid swamp | l | Ie
24 Swamp edge i {
foot || ae ea
Newnes Swamp
27 Mid swamp | i F r | ae ae
28 Swampedge F | le j 1 |
29 ‘Noodland = le ror 4 br bk b | L
Figure 2B. The pollen spectra from plant communities associated with swamps
within low frequency taxa. The Sample number (extreme left hand side) refers.
to the sample in Table 1
vegetation from which it came. For example, the
Myrtaceae pollen content (Figs. 2A, 3A), is lowest
from swamp sites, intermediate from the dry-land
communities bordering the swamps and highest from
the woodland and forest sites away from the swamps,
thus inferring a parallel approximate tree cover.
Swamp samples contain much higher pollen
frequencies of Restionaceae and/or Cyperaceae than
the dry-land sites, although both of these families
contain swamp and dry-land species. The species of
Restionaceae recorded in the vegetation (Appendix 1)
are found on wet and poorly drained soils and in damp
to wet heaths (PlantNet, 2007). Most of the species
of Cyperaceae, on the other hand, are found in fresh
water swamps and swampy areas (Sainty and Jacobs,
1981; PlantNet, 2007), although one dry-land species
is also recorded (Appendix 1). Thus high frequencies
Proc. Linn. Soc. N.S.W., 130, 2009
of Cyperaceae probably indicate swamps which are
more permanently waterlogged than swamps with
high frequencies of Restionaceae. Both Selaginella
and Gleichenia are found in wet places, on the edge
of swamps and streams (PlantNet, 2007).
The pollen of sclerophyllous shrub taxa (Figs
2B, 3B) are usually found sporadically and in very
low frequencies, indicating under-representation and
very localised distribution.
These findings are in accord with other studies
of surface pollen assemblages which indicate very
localised distribution of pollen (Dodson, 1983;
Kodela, 1990). Kershaw and Strickland (1990) found
that, in a 10 year pollen trapping experiment, most
pollen came from within 10 m of the trap.
These study sites are all contained within small
valleys where some barrier impedes drainage of the
117
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
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Figure 3A. Pollen spectra associat-
ed with dry-land plant communities
within major pollen groups. 1 The
sample number refers to the sam-
ple in Table 1. Codes for the vegeta-
tion map units are from Keith and
Benson (1988)
Proc. Linn. Soc. N.S.W., 130, 2009
118
J.M. CHALSON AND H.A. MARTIN
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Figure 3B. Pollen spectra associated with dry-land plant com-
munities within low frequency taxa. 1 The sample number
refers to the sample in Table 1. Codes for the vegetation map
units are from Keith and Benson (1988)
NN NY =| &
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stream and maintains the swamp (fora full description assemblage. While this may happen, it has been found
of the sites, see Chalson and Martin, this volume). It that very little pollen is transported into the site so
may be argued that pollen can be transported a long __ that the assemblage truly reflects the local vegetation
distance by a stream, to be deposited with the local (Chmura and Liu, 1990).
Proc. Linn. Soc. N.S.W., 130, 2009 119
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Table 2. Pollen type name on the pollen diagrams (Figs 2, 3) and the probable source in the vegetation.
Name on the pollen diagrams
2A and 3A
Podocarpus
Pinus
Angophora/Corymbia
Eucalyptus/Melaleuca
Melaleuca styphelioides
Leptospermum/Baeckea
Tristaniopsis
Unindetified Myrtaceae
Casuarinaceae
Poaceae
Restionaceae
Cyperaceae
Selaginella
Gleichenia
Other fern spores
Names on 2B and 3B
Grevillea acanthifolia
Grevillea
Hakea
Persoonia
Symphionema montanum
Banksia
Proteaceae
Acacia
Styphelia
Monotoca
Ericaceae
Rutaceae
Pimelea
Plantago
Haloragis
Asteraceae/Liguliflorae
Asteraceae/Tubulifiorae
Chenopodiaceae
120
Probable source in the vegetation and ecological inference.
From Plantnet (2007)
Probably Podocarpus spinulosus: sclerophyllous shrub/small tree
Pinus sp(p), Introduced: Pollen input from urban/forestry areas.
Species within the two genera: sclerophyll woodland
Species within the two genera : sclerophyll woodland/forest
Melaleuca styphelioides: moist stream bank habitat
Species within the two genera: ?mainly swamp communities
Tristaniopsis spp : moist habitats in sclerophyll communities
All pollen types not identifiable further
Casuarina, Allocasuarina sp(p): A distyla and A. nana in this study
Native and exotic species in the family: open situations, dryland
and swamp species
All species in the family: swamp and dry land species
All species in the family: swamp and dry land species
All species in the genus: damp sites, edge of swamp
Gleichenia sp(p): damp sites, edge of swamp
Other ferns: many possible species
G. acanthifolia: sclerophyllous understorey
Grevillea sp(p): sclerophyllous understorey
Hakea sp(p): sclerophyllous understorey
Persoonia sp(p): sclerophyllous understorey
S. montanum: heath or dry sclerophyll forest
Banksia sp(p): sclerophyllous understorey
Other taxa in the family sclerophyllous understorey
All species in the genus
Styphelia sp(p): sclerophyllous understorey
Monotoca sp(p): sclerophyllous understorey
Other taxa in the family: sclerophyllous understorey
All taxa in the family: sclerophyllous understorey
Pimelea sp(p): sclerophyllous understorey
Plantago sp(p): native and introduced herbs
Haloragis/Gonocarpus sp(p): Damp sites, sclerophyllous
understorey
Fenestrate-grained taxa in the subfamily Liguliflorae: herbs
Echinate-grained taxa in the subfam. Tubuliflorae: shrubs and herbs
Ruderals, salt tolerant
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
It is unfortunate that the Myrtaceae species
cannot be identified in most cases, since the vegetation
units are defined on their species of Myrtaceae. Most
Myrtaceae grains are small and thin-walled (Chalson,
1991; Chalson and Martin, 1995) and the preservation
may not be good enough to preserve this fine detail
which would distinguish the species. The result is
that there are large counts of unidentified Myrtaceae.
The alternate wetting and drying at the soil surface in
these sclerophyll forests are not ideal conditions for
pollen preservation.
The forests, woodlands and heaths defined by
Benson (1992), Keith and Benson (1988) and Benson
and Keith (1990) are structural units within one
major vegetation formation and share many species,
although the abundance of a particular species may
vary. The pollen assemblages cannot denote structure
of the vegetation and the floristics of these units are
too similar to allow any differentiation, especially
as the Myrtaceae pollen is so poorly preserved. For
practical purposes, the surface pollen assemblages can
denote major vegetation formations (Birks and Birks,
1980; Moore et al., 1991), more/less catchment tree
cover, swamp and/or adjacent dry-land environments
and local floral diversity.
ACKNOWLEDGEMENTS
We are indebted to the Joyce W. Vickery Research Fund
of the Linnean Society of NSW, the River Group Fund of
the Federation of University Women, and the Penrith Lakes
Development Corporation for financial assistance with this
project. Our thanks go to Dr. John Turner, the National Parks
and Wildlife Service of NSW and the Forestry Commission
of NSW for assistance. To the many friends, relatives and
colleagues who gave unstinting help and encouragement,
our heartfelt gratitude.
REFERENCES
Benson, D.H. (1992). The natural vegetation of the Penrith
1:100 000 map sheet. Cunninghamia 2(4), 502-662.
Benson, D.H. and Keith D.A. (1990). The natural
vegetation of the Wallerawang 1:100 000 map sheet.
Cunninghamia 2(2), 305-335
Birks, H.J.B. and Birks, H.H. (1980). ‘Quaternary
Palaeoecology’ (Edward Arnold, London) 289 pp.
Brown, C.A. (1960). ‘Palynological Techniques’ (C.A.
Brown, Baton Rouge) 188 pp.
Chalson, J.M. (1991). The late Quaternary vegetation
and climatic history of the Blue Mountains, NSW,
Australia. PhD Thesis, University of New South
Wales (unpubl.)
Proc. Linn. Soc. N.S.W., 130, 2009
Chalson, J.M. and Martin, H.A. (1995). The pollen
morphology of some co-occurring species of the
family Myrtaceae in the Sydney region. Proceedings
of the Linnean Society of New South Wales 115, 163-
Ii.
Chalson and Martin (this volume). A Holocene history of
the vegetation of the Blue Mountains, New South
Wales. Proceedings of the Linnean Society of New
South Wales 130, 77-109.
Chmura, G.L. and Liu, K-B. (1990). Pollen in the lower
Mississippi River. Review of Palaeobotany and
Palynology 64, 253-261.
Dodson, J.R. (1983). Modern pollen rain in southeastern
New South Wales, Australia. Review of Palaeobotany
and Palynology 38, 249-268.
Harden, G.J. (1992, 1993, 2000, 2002). ‘The Flora of
New South Wales, Vol. 3, Vol. 4, Vol.1 (revised
edition) and Vol. 2. (revised edition)’, respectively.
(University of New South Wales Press: Sydney).
Keith, D.A. and Benson, D.H. (1988). The natural
vegetation of the Katoomba 1:100 000 map sheet.
Cunninghamia 2(1), 107-145.
Kershaw, A.P. and Strickland, K.M. (1990). A 10 year
pollen trapping record from rainforest in northeastern
Queensland, Australia. Review of Palaeobotany and
Palynology 64, 281-288.
Kodela, P.G. (1990). Modern pollen rain from forest
communities on the Robertson Plateau, New South
Wales. Australian Journal of Botany 38, 1-24.
Moore, P.D., Webb, J.A. and Collison, M.E. (1991).
‘Pollen Analysis’. (Blackwell Scientific Publications,
Oxford).
PlantNet (2007). National Herbarium website (http://
plantNet.rbgsyd.nsw.gov.au) Accessed April 2007
Sainty, G.R. and Jacobs, S.W.L. (1981). “Waterplants of
New South Wales’ (Water Resources Commission of
N.S.W., Sydney) 550 pp.
121
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
APPENDIX. Species found in the vegetation on and around the swamps. Nomencla-
ture follows Harden (1992; 1993; 2000; 2002) and Plantnet (2006). Vegetation map
units are from Keith and Benson (1988) D, dominant. *, introduced species.
BURRALOW CREEK SWAMP
Species
BRYOPHYTES
Sphagnaceae
Sphagnum sp.
PTERIDOPHYTES AND ALLIES
Adiantaceae
Adiantum aethiopicum
Blechnaceae
Blechnum ambiguum
B. cartilaginum
Dennstaediaceae
Pteridium esculentum
Gleicheniaceae
Gleichenia dicarpa
G. microphylla
Osmundaceae
Todea barbara
Selaginellaceae
Selaginella uliginosa
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Platysace ericoides
P. lanceolata
P linearifolia
Xanthosia pilosa
Apocynaceae
Parsonsia straminea
Araliaceae
Polyscias sambucifolia
Asteraceae
Cassinia aculeata
C. aureonitens
Casuarinaceae
Allocasuarina nana
Ceratophyllaceae
Ceratophyllum demersum
Cunoniaceae
Callicoma serratifolia
Dilleniaceae
Hibbertia acicularis
H. bracteata
122
Wood-
land
10ag
Edge Mid
swamp swamp
28a 28a
+
+
+ +
+
+
+
+
+
+
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Elaeocarpaceae
Elaeocarpus reticulatus
Tetratheca thymifolia
Ericaceae
Epacris paludosa
E. pulchella
Leucopogon hookeri
Euphorbiaceae
Ampera xiphoclada
Phyllanthus hirtellus
Fabaceae, Faboideae
Bossiaea obcordata
Dillwynia floribunda
D. retorta
Gompholobium huegelii
Pultenaea tuberculata
Fabaceae, Mimosoideae
Acacia falciformis
A. myrtifolia
A. obtusata
A. ptychoclada
A. terminalis
Goodeniaceae
Dampiera stricta
Goodenia dimorpha
G. heterophylla
G. ovata
Lamiaceae
Prostanthera violacea
Lauraceae
Cassytha melantha
Lobeliaceae
Pratia purpurascens
Loganiaceae
Mitrasacme pilosa
Meliaceae
* Melia azedarach vat. australasica
Menyanthaceae
Villarsia exaltata
Myrsinaceae
Rapanea howittiana
Myrtacae
Angophora bakeri
A. costata
A. floribunda
Proc. Linn. Soc. N.S.W., 130, 2009
+
~ -
+
+
+ ~
+
+
+
+
+
+
D
123
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Baeckea linifolia
Corymbia eximia
Eucalyptus eugenioides
E. multicaulis
E. pauciflora
E. radiata
Kunzea capitata
Leptospermum polygalifolium
L. trinervium
Melaleuca linariifolia
Tristania neriifolia
Oleaceae
*Ligustrum sinense
Notelaea longifolia
Pittosporaceae
Billardiera scandens
Proteaceae
Banksia ericifolia
B. serrata
Hakea teretifolia
Lambertia formosa
Persoonia laurina
P. levis
P. linearis
P. mollis
P. oblongata
Petrophile pulchella
Ranunculaceae
Clematis aristata
Rhamnaceae
Cryptandra amara
Rutaceae
Eriostemon hispidulus
Sapindaceae
Dodonaea pinnata
D. triquetra
Stackhousiaceae
Stackhousia viminea
Thymelaeaceae
Pimelea ligustrina
Violaceae
Viola hederacea
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Baumea juncea
124
+
D
D
D
D
+
D
+
+
+
+
+
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Baumea sp. ar ar
Chorizandra sp. + zi
Eleocharis sphacelata + +
Lepidosperma longitudinale tt
Schoenus sp. a
Lomandraceae
Lomandra glauca a
L. longifolia +
Phormiaceae
Dianella caerulea =f
Restionaceae
Leptocarpus tenax ar
Baloskion fimbriatum a5
Smilacaceae
Smilax australis =
S. glyciphylla +f
WARRIMOO OVAL SWAMP Open Edge Mid
PTERIDOPHYTES AND ALLIES
Adiantaceae
Adiantum diaphanum a
Dennstaediaceae
Pteridium esculentum als ais
Gleicheniaceae
Gleichenia dicarpa + +
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Actinotus minor als
Platysace lanceolata a5 at
P linearifolia ar
Ericaceae
Brachyloma daphnoides a
Dracophyllum secundum +
Epacris paludosa a 7
Fabaceae, Faboideae
Bossiaea heterophylla “F a
*Cytisus scoparius ar
Daviesia ulicifolia ate +r
Dillwynia phylicoides a
Gompholobium huegelii + +
G. latifolium ae
Hovea linearis +
Proc. Linn. Soc. N.S.W., 130, 2009 125
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Mirbelia rubifolia
Fabaceae, Mimosoideae
Acacia falciformis
A. ptychoclada
A. rubida
A. terminalis
Goodeniaceae
Dampiera stricta
G. ovata
Lobeliaceae
Pratia purpurascens
Myrtacae
Angophora bakeri
Baeckea linifolia
Eucalyptus notabilis
E. pauciflora
E. radiata
Kunzea capitata
Leptospermum grandifolium
L. polygalifolium
L. trinervium
Polygalaceae
Comesperma defoliatum
C. ericinium
Proteaceae
Banksia ericifolia
B. oblongifolia
B. serrata
Grevillea laurifolia
G. mucronulata
G. phylicoides
Hakea salicifolia
Isopogon anethifolius
I. prostratus
Persoonia laurina
P. myrtilloides
P. pinifolia
Rutaceae
Boronia microphylla
Thymelaeaceae
Pimelea glauca
P. ligustrina
Violaceae
Viola hederacea
126
+
+++ +++ + 4+ 4+ 4+ 4+ 4+
+ + + +
~)
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Baumea juncea ar
Eleocharis sphacelata +
Juncaceae
Juncus remotiflorus ate
Lomandraceae
Lomanara filiformis ssp coriacea ae
L. longifolia ai
L. obliqua a5
Phormiaceae
Dianella caerulea +
Restionaceae
Leptocarpus tenax +f ap
NOTTS SWAMP Open -e/ « Mié
Species forest swamp
10ar 26a
PTERIDOPHYTES AND ALLIES
Dennstaediaceae
Pteridium esculentum a
Gleicheniaceae
Gleichenia dicarpa +
Selaginellaceae
Selaginella uliginosa +f
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Actinotus forsythii a
Platysace lanceolata +
P linearifolia +
Ericaceae
Epacris paludosa a
Lissanthe sapida ate
Euphorbiaceae
Poranthera microphylla ats
Fabaceae, Faboideae
Bossiaea heterophylla ste
Phyllota squarrosa 7°
Platylobium formosum +
Fabaceae, Mimosoideae
Acacia melanoxylon al
A. obtusata =i
A. obtusifolia a
A. stricta a
Proc. Linn. Soc. N.S.W., 130, 2009 WF
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Myrtacae
Eucalyptus aggregata
E. dives
E. ligustrina
E. pauciflora
E. piperita
E. sclerophylla
Kunzea capitata
Leptospermum juniperinum
Proteaceae
Banksia oblongifolia
B. serrata
Grevillea phylicoides
Hakea teretifolia
Isopogon prostratus
Persoonia laurina
P. linearis
Petrophile pedunculata
Rutaceae
Boronia microphylla
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Baumea rubiginosa
Carex sp.
Gahnia sp.
Iridaceae
Patersonia sericea
Juncaceae
Juncus remotiformis
Phormiaceae
Dianella caerulea
Poaceae
Entolasia marginata
Poa sp.
Restionaceae
Baloskion australe
Leptocarpus tenax (Labill.)
INGAR SWAMP
Species
GSeoeagvs
+ ++ + + + + +
Open
forest
10ar
Wood- 6cTall Edge Mid
PTERIDOPHYTES AND ALLIES
Adiantaceae
Adiantum aethiopicum
128
land open swamp swamp
10ag forest 26a 26a
+
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Dennstaediaceae
Pteridium esculentum
Dicksoniaceae
Calochlaena dubia
Gleicheniaceae
Gleichenia dicarpa
G. microphylla
Osmundaceae
Todea barbara
Selaginellaceae
Selaginella uliginosa
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Actinotus forsythii
Platysace lanceolata
P. linearifolia
Casuarinaceae
Allocasuarina distyla
Cunoniaceae
Bauera rubioides
Callicoma serratifolia
Ceratopetalum apetalum
Dilleniaceae
Hibbertia acicularis
Elaeocarpaceae
Elaeocarpus reticulatus
Ericaceae
Brachyloma daphnoides
Dracophyllum secundum
Epacris paludosa
Leucopogon esquamatus
L. hookeri
L. lanceolatus
Lissanthe sapida
Euphorbiaceae
Ampera xiphoclada
Fabaceae, Faboideae
Bossiaea heterophylla
B. obcordata
Daviesis alata
D. ulicifolia
Dillwynia philicoides
D. retorta
Glycine clandestina
Hovea linearis
Proc. Linn. Soc. N.S.W., 130, 2009
+
+
129
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Phyllota phylicoides
P. squarrosa
Platylobium formosum
Pultenaea divaricata
P. flexilis
P. incurvata
P. tuberculata
Fabaceae, Mimosoideae
Acacia echinula
. melanoxylon
. obliquinervia
. obtusata
. obtusifolia
. stricta
DS TX JX EX Bx SS
. suaveolens
Goodeniaceae
Dampiera stricta
Goodenia bellidifolia
G. dimorpha
G. ovata
Haloragaceae
Gonocarpus chinensis ssp verrucosus
G. longifolius
Myrtacae
Angophora bakeri
Backhousia myrtifolia
Baeckea diosmifolia
Corymbia eximia
Eucalyptus agglomerata
E. dalrympleana
E. dives
E. obliqua
E. oreades
E. pauciflora
E. radiata
E. sieberi
Kunzea capitata
Leptospermum grandifolium
L. juniperinum
L. polygalifolium
L. scoparium
L. trinervium
Melaleuca linariifolia
Syncarpia glomulifera
130
+ + + +
+
+
+
+
+
- +
+ +
+
+
+
+
+
+
+ +
+
+
D
+
D
+
+
+
D
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Proteaceae
Banksia ericifolia
B. oblongifolia 3 af
B. serrata + +
Grevillea aspleniifolia +
G. laurifolia
G. phylicoides =F
Hakea propinqua
H. sericea or
H. teretifolia ai af
Isopogon prostratus ai ar
Lambertia formosa +
Lomatia myricoides
Persoonia acerosa ate
P. laurina +
P. levis ate
P. linearis =F +r
P. pinifolia 1°
Petrophile pedunculata als
Ranunculaceae
Clematis aristata
Rhamnaceae
Cryptandra amara
Rutaceae
Boronia microphylla a
Thymelaeaceae
Pimelea ligustrina
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Baumea rubiginosa
Carex sp. + aie
Chorizandra cymbaria
Eleocharis sphacelata cle
Gahnia sieberana a
Gahnia sp.
Gymnoschoenus sphaerocephalus
Lepidosperma longitudinale
Iridaceae
Patersonia sericea ar
Juncaceae
Juncus remotiformis
Luzuriagaceae
Eustrephus latifolius
Phormiaceae
Dianella caerulea ats
Proc. Linn. Soc. N.S.W., 130, 2009
‘31
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Poaceae
Entolasia marginata
Poa sp.
Restionaceae
Baloskion australe
Empodisma minus
Leptocarpus tenax
Smilacaceae
Smilax australis
KINGS TABLELAND SWAMP
Species
PTERIDOPHYTES AND ALLIES
Dennstaediaceae
Pteridium esculentum
Gleicheniaceae
Gleichenia dicarpa
GYMNOSPERMS
Cupressaceae
Callitris muelleri
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Actinotus forsythii
Platysace lanceolata
Casuarinaceae
Allocasuarina distyla
Allocasuarina nana
Ericaceae
Dracophyllum secundum
Epacris paludosa
Fabaceae, Faboideae
Bossiaea heterophylla
Daviesia alata
D. ulicifolia
Hovea linearis
Phyllota squarrosa
Pultenaea divaricata
Fabaceae, Mimosoideae
Acacia obtusata
A. stricta
A. suaveolens
A. terminalis
132
Open
forest
91
+
+
D
D D
+
Open Edge Mid
heath swamp swamp
GE 26a 26a
+
Proc. Linn. Soc. N.S.W., 130, 2009
Myrtacae
Corymbia eximia
C. gummifera
Eucalyptus deanei
E. oblonga
E. pauciflora
E. piperita
E. sclerophylla
E. stellulata
E. stricta
Kunzea capitata
K. ericoides
Leptospermum grandifolium
L. juniperinum
L. polygalifolium
Olacaceae
Olax stricta
Proteaceae
Banksia ericifolia
B. oblongifolia
B. serrata
B. spinulosa
Grevillea phylicoides
Hakea dactyloides
H. salicifolia
H. sericea
Isopogon anemonifolius
I. prostratus
Lomatia silaifolia
Persoonia laurina
Petrophile pedunculata
Thymelaeaceae
Pimelea ligustrina
J.M. CHALSON AND H.A. MARTIN
ANGIOSPERMS, MONOCOTYLEDONS
Lomandraceae
Lomandra glauca
KATOOMBA SWAMP
Species
BRYOPHYTES
Dawsoniineae
Dawsonia sp.
Open
forest
Upper 91
Open
forest
Lower 91
Edge
swamp
26a
Proc. Linn. Soc. N.S.W., 130, 2009
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
PTERIDOPHYTES AND ALLIES
Blechnaceae
Blechnum cartilaginum
Dennstaediaceae
Pteridium esculentum
Gleicheniaceae
Gleichenia dicarpa
Lycopodiaceae
Lycopodium deuterodensum
ANGIOSPERMS, DICOTYLEDONS
Araliaceae
Polyscias sambucifolia
Asteraceae
Arrhenechthites mixta
Bracteantha bracteata
Cunoniaceae
Callicoma serratifolia
Ericaceae
Epacris paludosa
Fabaceae, Faboideae
Bossiaea rhombifolia
Daviesia latifolia
Fabaceae, Mimosoideae
Acacia obtusata
A. suaveolens
Myrtacae
Callistemon citrinus
Eucalyptus obliqua
E. oblonga
E. sclerophylla
E. squamosa
Kunzea capitata
K. ericoides
Leptospermum polygalifolium
L. trinervium
Oleaceae
*Ligustrum sinense
Polygonaceae
*Acetosella vulgaris
*Rumex obtusifolius
Proteaceae
Banksia spinulosa
Grevillea mucronata
Isopogon prostratus
Lomatia myricoides
134
+
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Persoonia laurina
Petrophile pedunculata
Rutaceae
Boronia microphylla
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Caustis flexuosa
Juncaceae
Juncus remotiformis
Lomandraceae
Lomandra obliqua
Phormiaceae
Dianella caerulea ae
Poaceae
Poa sp.
NEWNES SWAMP Open
Species rey
91
PTERIDOPHYTES AND ALLIES
Blechnaceae
Blechnum cartilaginum ata
Dennstaediaceae
Pteridium esculentum +
Gleicheniaceae
Gleichenia dicarpa a”
ANGIOSPERMS, DICOTYLEDONS
Apiaceae
Platysace lanceolata F
Asteraceae
Arrhenechthites mixta ate
Helichrysum scorpioides aL
Olearia sp. aff. chrysophyplla ate
Casuarinaceae
Allocasuarina nana
Dilleniaceae
Hibbertia dentata ay
Ericaceae
Brachyloma daphnoides
Epacris obtusifolia
E. paludosa ae
Lissanthe sapida fe
Monotoca scoparia
Proc. Linn. Soc. N.S.W., 130, 2009
Wood-
land
lla
Edge
swamp
20a
+
Mid
swamp
20a
jam
ore)
CA
MODERN POLLEN DEPOSITION IN THE BLUE MOUNTAINS
Euphorbiaceae
Ampera xiphoclada
Fabaceae, Faboideae
Daviesis corymbosa
D. ulicifolia
Gompholobium grandiflorum
G. latifolium
Phyllota phylicoides
P. squarrosa
Platylobium formosum
Fabaceae, Mimosoideae
Acacia elata
A. linifolia
A. longifolia
A. melanoxylon
A. suaveolens
Goodeniaceae
Dampiera stricta
Myrtacae
Baeckea diosmifolia
Eucalyptus acmenoides
E. aggregata
E. deanei
E. notabilis
E. oreades
E. racemosa
E. sclerophylla
Kunzea capitata
Leptospermum juniperinum
L. polygalifolium
Proteaceae
Banksia spinulosa
Grevillea acanthifolia
G. aspleniifolia
G. phylicoides
Hakea salicifolia
H. teretifolia
Petrophile pedunculata
Ranunculaceae
Clematis aristata
Rhamnaceae
Cryptandra amara
Rutaceae
Boronia microphylla
136
+ ++ 4
SGeevwes
o
+
+
+ +
D
+ +
+ D
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Santalaceae
Exocarpos strictus +
Thymelaeaceae
Pimelea glauca oF
P. ligustrina a
ANGIOSPERMS, MONOCOTYLEDONS
Cyperaceae
Lepidosperma laterale +
Iridaceae
Patersonia sericea + +
Juncaceae
Juncus remotiformis a
Lomandraceae
Lomanadra filiformis ssp coriacea tr or
L. filiformis ssp filiformis ats
L. glauca Ewart 4
Phormiaceae
Dianella caerulea +
Poaceae
Entolasia marginata
Poa sp. +
Restionaceae
Empodisma minus
Leptocarpus tenax vi 1
Proc. Linn. Soc. N.S.W., 130, 2009
onounibre
aonkvers ee iy cOVNA, cotbint
aR, ae rancuelNR,
hewel oomnly,
eWlenincen: — ymooainnt
eulrreng BUSH
PANTY BIE
a snethe ew
rons)
Silurian Rhynchonellide Brachiopods from Yass, New South
Wales
Desmonp L. Strusz
Department of Earth and Marine Sciences, Research School of Earth Sciences, Australian National
University, Canberra, Australia 0200 (dstrusz@ems.anu.edu.au), and Research Associate, Australian
Museum, 6 College Street, Sydney, NSW 2010.
Strusz, D.L. (2009). Silurian rhynchonellide brachiopods from Yass, New South Wales. Proceedings of
the Linnean Society of New South Wales 130, 139-146.
Rhynchonellide brachiopods are rare in the Silurian sequence at Yass. In this paper two species are
described, one new species Agarhynchus australe being abundant at just one locality in the late Wenlock or
earliest Ludlow Yass Formation. The other species, tentatively assigned to Tuvaerhynchus, is known from
only a few specimens of late Wenlock to Ludfordian age.
Manuscript received 19 October 2008, accepted for publication 21 January 2009.
KEYWORDS: Agarhyncha australe, Ludlow, rhynchonellide, Silurian, Tuvaerhynchus, Wenlock, Yass
INTRODUCTION
Rhynchonellide brachiopods were recognised
in early accounts of the stratigraphy of the Yass
Syncline, but none has ever been described. Jenkins
(1879, p. 26) recorded Rhynchonella from what
(using modern terminology) was probably the basal
Bowspring Limestone at locality GOUS7, and Mitchell
(1887, p. 1201) listed the same genus from pebbles
in the Sharpeningstone Conglomerate at Bowning
(two specimens, described in this paper). Shearsby
(1912, pp. 110-112) in his more detailed account of
the succession north of Yass then noted the presence
of possible Rhynchotreta and Camarotoechia at two
localities, one within the Douro Volcanics, the other
in the Yass Formation. The latter is in the same area
along Derringullen Creek from which both of the
species described in this paper were collected by Dr
R.S. Nicoll and myself in 1982. However, other than
at that locality, rhynchonellides are rare (only six
usable specimens) in the Yass sequence.
Only two taxa can be recognised. The first,
Agarhyncha australe n. sp., occurs at only the
one locality (on Derringullen Creek), just below
the Cliftonwood Limestone Member of the Yass
Formation, but is there in some numbers. Agarhyncha
Havliéek, 1982, is otherwise known from the Wenlock
and Ludlow of the Czech Republic. The other Yass
rhynchonellide occurs in very low numbers at a few
localities from the Yass Formation to the Yarwood
Siltstone Member of the Black Bog Shale, and in
pebbles in the Sharpeningstone Conglomerate. It
is tentatively referred to the genus 7uvaerhynchus
Kul’kov, 1985, from the Wenlock of Tuva. This
raises some problems concerning provinciality which
cannot be properly assessed until better material from
Yass becomes available, enabling more confident
identification.
For a diagrammatic representation of Yass
stratigraphy and ages, refer to Strusz (2002, fig.
1). Localities are detailed in that publication, with
additions in Strusz (2003, 2005).
SYSTEMATIC PALAEONTOLOGY
Classification
The classification followed is that of Savage et
al. (2002).
Measurements and symbols
All linear measurements are in millimetres,
and unless otherwise specified are as defined by
Williams and Brunton (1997); the following symbols
are used for these measurements:
Ls, Ws, Ts — maximum shell length, width,
thickness.
Wh — hinge width.
L(Wmax) — length to widest part of shell.
SILURIAN BRACHIOPODS FROM YASS
Repositories
The repositories for the specimens studied are
shown by the following prefixes to their catalogue
numbers:
AMF - macrofossil
Museum, Sydney.
ANU - Department of Earth and Marine Sciences
(Research School of Earth Sciences),
Australian National University, Canberra.
CPC - Commonwealth Palaeontological
Collection, Geoscience Australia, Canberra.
collection, Australian
Phylum BRACHIOPODA
Class RHYNCHONELLATA Williams, Carlson,
Brunton, Holmer and Popov, 1996
Order RHYNCHONELLIDA Kuhn, 1949
Superfamily RHY NCHOTREMATOIDEA
Schuchert, 1913
Family TRIGONIRHYNCHIIDAE Schmidt, 1965
Subfamily TRIGONIRHYNCHINAE Schmidt,
1965
Genus Agarhyncha Havliéek, 1982
Type species
Terebratula famula Barrande, 1847, by original
designation; Ludlow, Bohemia.
Diagnosis
Subpentagonal to subcircular outline;
biconvex to globose profile. Beak suberect to erect;
foramen with minute deltidial plates. Fold and sulcus
well defined, broad, anterior commissure uniplicate;
tongue rectangular, serrate. Costae coarse, rounded,
simple, but umbones smooth. Dental plates very short.
Dorsal median septum thin; septalium with cover
plate anteriorly; crura close to septum posteriorly
(Savage p. 1052 in Savage et al. 2002).
Agarhyncha australe sp. nov.
Figs 1-5, Table 1
Diagnosis
Relatively large biconvex species of
Agarhyncha with smooth non-sulcate umbones,
sulcus often weak anteriorly, ribs only moderately
developed, medially concave dental plates, impressed
ventral muscle field, raduliform crura, long dorsal
median septum.
Material
Holotype CPC39529, paratypes CPC39530-
39592, all from locality GOU49.
140
Horizon
Topmost O’Briens Creek Member, Yass
Formation.
Age
Probably Homerian (late Wenlock), possibly
earliest Gorstian (early Ludlow).
Description
Juvenile shells (taken as Ws <6.0 mm - see
Fig. 4b) lenticular, biconvex to ventribiconvex,
elongate lacriform to lozenge-shaped (mean juvenile
Ls/Ws 1.10, mostly 1.0-1.2), generally relatively
thin (mean juvenile Ts/Ws 0.43, mostly 0.35-
0.50). Adult shells (Ws >6 mm) subtriangular to
subpentagonal, biconvex to slightly ventribiconvex,
largest shells globose (mean adult Ts/Ws 0.55, max.
0.85). Maximum observed width 12.2 mm; length
about equal to width (mean adult Ls/Ws 1.01, mostly
0.9 - 1.1). Dorsal fold and ventral sulcus appear at
lengths of 3-4 mm, generally shallow, but variably
developed anteriorly in larger shells; tongue when
developed trapezoidal. Ventral beak suberect, sharp
(especially in juveniles), usually small but in some
shells extended posteriorly. Foramen mesothyrid
(Fig. lh), delthyrium wide, deltidial plates narrow,
disjunct. Umbones smooth, ribs appearing at Ls from
2.5 to 5 mm, initially faint. Ribs anteriorly rounded-
angular, simple, generally low (especially laterally);
margins of sulcus defined by pair of relatively well
developed ribs, sulcus contains 1-3 ribs (2-4 on fold);
2-5 ribs on each flank.
Shell generally thin-walled. Dental plates short,
upright to gently convergent ventrally, somewhat
concave medially. Ventral muscle field elongate,
moderately impressed into slightly medially thickened
shell, may be divided by very low myophragm.
Dorsal median septum long (at least Ls/2), posteriorly
supports small V-shaped septalium (Fig. 3) which is
open posteriorly, covered mid-length to anteriorly (see
Fig. 2, especially CPC39544, sections 1.2 to 1.8 mm).
Outer hinge plates wide, flat in narrow zones between
crural bases and inner socket ridges, moderately thick
medially and generally strongly thickened beneath
sockets. Sockets widely divergent, large; inner socket
ridges robust, outer socket ridges merged with valve
walls. Crural bases strong, triangular; crura calciform,
curved somewhat towards ventral valve. No cardinal
process.
Remarks
This form differs from leiorhynchids in its
generally thin-walled shell which is mostly not
globose, in its only moderately impressed ventral
Proc. Linn. Soc. N.S.W., 130, 2009
DL. STRUSZ
Figure 1. Agarhyncha australe; a-g, growth series of paratype shells in dorsal aspect, CPC39535, 39536,
39537, 39540, 39539, 39541, 39538; h-l, holotype CPC39529 in dorsal, lateral, ventral, posterior and an-
terior aspects; m-p, paratype CPC39532 in dorsal, lateral, ventral and anterior aspects, a partly decorti-
cated relatively wide shell with anteriorly well developed fold; q-s, paratype CPC39533 in dorsal, lateral
and ventral aspects, a posteriorly decorticated shell with low convexity, few subdued ribs; t, paratype
CPC39543, a large shell in ventral aspect, with 4 anteriorly strong ribs in sulcus; u-w, paratype CPC
39534 in dorsal, lateral and ventral aspects, a posteriorly decorticated large shell showing local crush-
ing, presumably before lithification of the enclosing sediment; x, paratype CPC 39542, a large relatively
wide shell in ventral aspect; y, paratype CPC39592, a dorsal internal mould (see Fig. 3). All x4, scale
bar 5 mm. Locality GOU49, Yass Formation, O’Briens Creek Member immediately below Cliftonwood
Limestone; probably Late Wenlock.
Proc. Linn. Soc. N.S.W., 130, 2009 [4]
SILURIAN BRACHIOPODS FROM YASS
CPC39544
OOO OL
Yi ie *
U2 14
CPC39545
Figure 2. Agarhyncha australe; selected serial sections of paratypes CPC39544, 39545; distances from
posterior ends in millimetres. Scale bar 2 mm.
2mm
Figure 3. Agarhyncha australe; paratype dor-
sal internal mould CPC39592 enlarged to show
septalium and long but low median septum. Scale
bar 2 mm.
muscle field, and a cover plate on the septalium. From
rhynchotrematids it differs in its smooth umbones,
distinct dental plates and lack of a cardinal process. It
shares important features with the Trigonirhynchiidae.
Among trigonirhynchiids Astua Havlitek, 1992
(Lochkovian, Bohemia and central Asia) differs in
stronger ribs, fold and sulcus, an emarginate anterior
commissure, and internally in lacking a cover-plate on
the septalium. Oxypleurorhynchia Plodowski, 1973
(Pridoli, Carnic Alps) is dorsibiconvex, with coarse
ribs and pronounced fold and sulcus extending from
the umbones. Virginiata Amsden, 1968 (Llandovery
to Ludlow, N. America, China and Siberia) lacks fold
and sulcus, but its ribs extend from the beaks; it also
differs in being more elongate, having a posterior
cover-plate on the very small septalium, robust
cardinalia, and a short dorsal median septum. The
new species is referred to the Bohemian Wenlock
to Ludlow genus Agarhyncha on the basis of its
142
Ls = 1.08Ws
Ts =
O.48\/s
Ws mm
) ead (CRs
4 8 12
Figure 4. Agarhyncha australe; a, length (Ls) and
thickness (Ts) plotted against width (Ws). The di-
vergence from the overall means at widths above
6 mm is just noticeable on these plots; b, thick-
ness (Ts) plotted against width (Ws) on log-normal
coordinates; in this plot the change in growth pa-
rameters at a width of about 6 mm is quite clear.
smooth umbones, short dental plates, and medially to
anteriorly covered septalium.
The Ludlow-age type species, Agarhyncha
famula (Barrande, 1847) is smaller (Ws to c. 9.6
Proc. Linn. Soc. N.S.W., 130, 2009
D.L.
8= ~—L(Wmax) mm
LiWmax) = 0.61Ls
Figure 5. Agarhyncha australe; plot of length to
greatest width (L(Wmax)) against length (Ls),
showing only weak variability.
mm), more globose, with in some cases anteriorly
truncated margins, often posteriorly elongate ventral
beak, stronger ribs which may be flattened and
grooved marginally, shallower sulcus, ribs at least
faintly developed umbonally, high dorsal median
septum, and rod-like crura. The Wenlock species A.
agason Havliéek in Havliéek and Storch, 1990 is
of comparable size and outline, but has more and
stronger, more angular ribs, especially medially.
The other Bohemian Ludlow species, A. chuchlensis
Havliéek in Havliéek and Storch, 1990 is wider
(Ls/Ws_ 0.83-0.95), with generally subpentagonal
outline, low, rounded beak, weakly ribbed umbones,
anteriorly well developed fold and sulcus, more ribs,
ventral muscle field not impressed but dorsal adductor
field with fine lateral bounding ridges, rod-like crura,
and somewhat shorter dorsal median septum. None of
the Bohemian species shows medially concave dental
plates.
Family ORTHORHYNCHULIDAE Cooper, 1956
Genus Tuvaerhynchus Kulkov, 1985
Type species
Tuvaerhynchus khalfini Kul’ kov in Kul’kov et
al., 1985, by original designation; Wenlock, Tuva.
Diagnosis
Small with subpentagonal to subrectangular
outline and dorsibiconvex profile. Beak suberect;
delthyrium with disjunct deltidial plates. Fold and
sulcus strong, narrow, well defined, from umbones;
anterior commissure uniplicate; tongue high,
trapezoid, dentate. Costae numerous, simple, angular.
Dental plates short, vertical, close to valve wall.
Septalium short, wide; hinge plates concave, slope
medially; cardinal process septiform, thin; crura short,
curved sharply ventrally (Savage p. 1081 in Savage et
al. 2002).
Proc. Linn. Soc. N.S.W., 130, 2009
STRUSZ
Tuvaerhynchus? sp.
Fig. 6, Table 2
Material
Yass Formation: GOU47, CPC 39595 and
1 very uncertain fragment; GOU49, CPC 39596.
Barrandella Shale Member, Silverdale Formation:
GOU2a, CPC 39593. Lower Black Bog Shale: KF,
ANU46537. Yarwood Siltstone Member, Black Bog
Shale: GOU28, CPC 39594. Horizon uncertain:
Bowning, “Upper Conglomerate”, Mitchell Collection
AMEF28588, 133959 - presumably (following
Mitchell 1887) from pebbles in the Sharpeningstone
Conglomerate, derived from an older horizon.
Stratigraphic distribution
Yass Formation to Yarwood Siltstone Member,
Black Bog Shale
Age
Late Wenlock? to early Ludfordian
Description
Available specimens are few, and mostly
poorly preserved. Best are a steinkern from the
Mitchell Collection, and a small shell from GOU47.
Both are dorsibiconvex, with rounded outline; the
small shell (CPC 39595, Ws 6.2 mm) is longer than
wide (Ls/Ws ca 1.16), the steinkern (AMF28588, Ws
ca 12 mm) transversely oval (Ls/Ws ca 0.8). They are
globose - in both cases Ts/Ls is about 0.7. They are
strongly ribbed, the ribs starting at the beaks. CPC
39595 has a shallow ventral sulcus with 3 ribs, there
being 5 ribs on each flank. AMF28588 has a well
developed fold and sulcus, forming a high trapezoidal
tongue anteriorly; the sulcus contains 3 ribs, the flanks
5 ribs each, and the fold is formed of 2 ribs which
split once. The other Mitchell Collection specimen,
AMF133959, is an incomplete flattened internal
mould with 4 ribs in the sulcus, 6 on each flank. Inter-
rib furrows extend as short marginal spines. None of
the specimens shows clear details of the ventral beak,
and so the presence and nature of a delthyrium cannot
be demonstrated.
Large teeth are supported by fairly short but
distinct dental plates which are somewhat convergent
towards the valve floor. Details of the ventral muscle
field are not known. Dorsal median septum long, low,
fine, continuous with linear cardinal process which
arises from a shallow septalium which is either sessile
or nearly so. Crural bases robust, crura unknown.
Discussion
Among Silurian rhynchonellides, the general
SILURIAN BRACHIOPODS FROM YASS
Ls Ld Ws
CEE39529"; OF me SION OF
CPC39530 5975. Se, (6.0,
CPC39534 9.0 - 8.9
CPC39538 Be) a SM)
CPC39543 9:9 - 9.6
Ts L(Wmax) Ls/Ws _ Ts/Ws
3.3 3.4 0.97 0.52
39) 3.3 0.98 0.65
5.0 4} On 0.56
es) 4.0 1.18 0.50
- De) 1.03 -
Table 1: Agarhyncha australe: dimensions in mm and proportions of holotype (*) and selected para-
types. Measurements in italics are best estimates for damaged specimens.
Ls Ws Ts
AMEF28588 DT NZ FO)
CPC39593 NOS Qn 8
CPC39595 TO re
ANU46537 iS Sai) -
L(Wmax) Ls/Ws _ Ts/Ws
4.8 0.81 0.58
55 HT 0.56
4.0 LAG 0.82
55) 0.94 -
Table 2. Tuvaerhynchus? sp.: dimensions in mm and proportions of selected specimens. Measurements
in italics are best estimates for damaged specimens.
shell form and strong simple ribbing of this form,
coupled with distinct but short dental plates and a
linear cardinal process on a sessile or near-sessile
septalium, points to the Orthorhynchulidae (whose
genera are also united by possessing an open or near-
open delthyrium). Orthorhynchula Hall and Clarke,
1893, has dental plates fused to the valve walls, and
a low fold. The Tasmanian Ordovician Tasmanella
Laurie, 1991, has a high fold, but differs in its fused
dental plates, and a short, high dorsal median septum
supporting a raised septalium. Twvaerhynchus is
closest morphologically, but in the absence of details
of delthyrium, deltidial plates, and crura, generic
identity cannot be certain. In the absence of that
certainty, palaeobiogeographic speculation on this
possible link between the Tuvaella and Retziella
Faunas of Rong et al. (1995), and thus the Mongolo-
Okhotsk and Sino-Australian Provinces of the
Uralian-Cordilleran Region, is pointless.
ACKNOWLEDGEMENTS
I thank Jan Percival and Norman Savage for
reviewing this paper, and Prof. Brian Kennett, Director of
the ANU Research School of Earth Sciences, for continuing
provision of facilities in the Department of Earth and
Marine Sciences within that School. Serial sectioning was
made possible by the loan of a Croft Parallel Grinder from
the Geological Survey of New South Wales. Photography
by H.M. Doyle (formerly of Geoscience Australia) and the
author.
144
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Amsden, T. W. (1968). Articulate brachiopods of the St. Clair
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Barrande, J. (1847). Uber die Brachiopoden der silurischen
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Cooper, G.A. (1956). Chazyan and related brachiopods.
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Havlicek, V. (1992). New Lower Devonian (Lochkovian-
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Havliéek, V. and Storch, P. (1990). Silurian brachiopods
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Jenkins, C. (1879). On the geology of Yass Plains.
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Sweizerbart’sche Verlagsbuchhandlung, Stuttgart).
Proc. Linn. Soc. N.S.W., 130, 2009
DL. STRUSZ
Figure 6. Tuvaerhynchus? sp.; a-e, CPC39593, a slightly crushed lenticular shell in dorsal, lateral, ven-
tral, posterior and anterior aspects (locality GOU2a, Barrandella Shale Member, Silverdale Fm, late
Gorstian); f-j, CPC39595, a shell in dorsal, lateral, ventral, posterior and anterior aspects, the dorsal
umbo worn and revealing the median septum (locality GOU47, topmost O’Briens Creek Member, Yass
Fm, probably Late Wenlock); k-o, AMF28588, a wide and very globose steinkern in dorsal, lateral, ven-
tral, posterior and anterior aspects, the ventral beak broken (Mitchell Collection, from pebble in Sharp-
eningstone Creek Conglomerate); p, AMF133959, somewhat crushed ventral internal mould (source as
AMEF728588). All x4, scale bar 5 mm.
ee
Bl
A
Proc. Linn. Soc. N.S.W., 130, 2009
SILURIAN BRACHIOPODS FROM YASS
Kul’kov, N.P., Vladimirskaya Ye.V. and Rybkina N.L.
(1985). Brakhiopody i biostragrafiya verkhnego
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Laurie, J.R. (1991). Articulate brachiopods from the
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Rong J.-Y., Boucot, A.J., Su, Y.-Z. and Strusz, D.L.,1995.
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1911, 106-119.
Strusz, D.L. (2002). Brachiopods of the Orders Protorthida
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146
Proc. Linn. Soc. N.S.W., 130, 2009
Cortinarius Fr. Subgenus Cortinarius in Australia
A.E.Woop:
School of Biological, Earth and Environmental Sciences, University of New South Wales, UNSW Sydney,
NSW, 2052, Australia.
Wood, A.E. (2009). Cortinarius Fr. subgenus Cortinarius in Australia. Proceedings of the Linnean Society
of New South Wales 130, 147-155.
Three new species within Cortinarius subgenus Cortinarius from Australia are described, each belonging
near a different species, but differing significantly from the type variety in all cases. They represent distinct
species — C. jenolanensis, C. kioloensis and C. hallowellensis.
Manuscript received 15 October 2008, accepted for publication 4 February 2009
Keywords: agarics, Cortinarius, distribution, mushrooms, new varieties, toadstools
INTRODUCTION
Cortinarius subgenus Cortinarius is characterised
by the presence of fleshy carpophores, with a cap
that is frequently squamulose, large conspicuous
cheilocystidia and vacuolar, mostly violet, pigments.
The spores show both a suprapilar plage, usually
flattened and often more or less smooth.
There have been scattered records of this
subgenus, particularly C. violaceus from Australia.
This species was reported from Victoria by Cooke
(1892) and this report was carried forward by
McAlpine (1895) and Brittlebank (1940). Cleland
(1933, 1934) did not record the species, nor did
Grgurinovic (1997) record it from South Australia.
Shepherd and Totterdell (1988) recorded the species
from the Australian Capital Territory, New South
Wales and Victoria. Young (1994) also recorded the
species from New South Wales and Victoria. This
species was also recorded from Western Australia
by Griffiths (1985), Hilton (1988) and Syme (1992)
and more recently was fully described by Bougher
and Syme (1998). All these records are for C.
violaceus, in some cases with uncertainty being
expressed as to whether the collections are identical
with the European species. Recently a new species,
Cortinarius austroviolaceus has been described from
Tasmania by Gasparini (2001).
There have been some recent studies on C.
violaceus in Europe and now two species are widely
recognised, C. violaceus and C. hercynicus (Brandrud
1983; Brandrud et al. 1989-1998). The study by Moser
(1986) of some collections from the SW-Pacific area
has added four more species to the subgenus C.
atroviolaceus, C. subcalyptrosporus, C. atrolazulinus
and C. paraviolaceus. In view of the diversity of taxa
of the subgenus in the SW Pacific, the suggestion
has been made that they represent the descendants
of a Gondwanan species of possibly ancient origin
(Gasparini, 2001). However the subgenus has not
been reported from Tierra del Fuego (Horak 1979) or
in other areas of South America (Moser and Horak
1975). Cortinarius violaceus s.s. Montagne, (from
Chile, see Horak, 1979) is a different, unrelated
species, Cortinarius gayii Horak (see Horak, 1979, p.
396 with full description).
There has been considerable discussion over
many years as to whether Cortinarius violaceus is
a single species in Europe or whether several taxa
at some close level (species, subspecies or variety)
are involved. Some claim that over a large number
of collections, a continuous variation can be found
between the two main forms. However many now
recognise two distinct forms, though the level at
which they should be considered is also disputed.
The view taken here (following Moser (1983), Horak
(2005), Breitenbach & Kranzlin (2000) and Knudsen
&Vesterholt (2008)) is to recognise two separate
species from Europe as follows :
Cortinarius violaceus with spores (12)13-16(17)
x 7-8(8.5) um, elliptic to amygdaliform, verrucose,
cap mostly 6-14 cm, under deciduous woods;
Cortinarius hercynicus with spores (12)13-
16(17) x 7-8(8.5) um, broadly ellipsoid to subglobose,
strongly verrucose, cap mostly 5-10 cm, under
coniferous woods (spruce, pine, sometimes mixed
woods).
CORTINARIUS Fr. SUBGENUS CORTINARIUS IN AUSTRALIA
Most records are only from the latter part of the
twentieth century (May and Wood, 1997). The records
are probably accurate because of the distinctive
characteristics of Cortinarius violaceus s.1., but they
give no information as to which of the currently
reported species are intended. Later records indicate
that the subgenus is widespread throughout most of
Australia, but that it is not collected frequently.
Studies of DNA sequences of various species
of Cortinarius concluded that there were grounds
for considering the creation of two separate genera
(Hoiland and Holst-Jensen, 2000). A later study
of DNA sequences for a large range of Cortinarius
species (Garnica et al, 2005) supported the Cortinarius
clade, without any further additions of any closely
related groups or species. Bougher and Syme (1998)
used the epithet C. violaceus with some reservations
for their local collections. Chambers et al. (1999)
compared DNA from New South Wales material
with reported sequences from Northern Hemisphere
collections of Cortinarius violaceus, and reported that
the local material while close, belonged to a different
taxon and noted ‘a careful revision of Australian
Cortinarius violaceus collections is clearly required’.
Unfortunately, voucher material of these collections
has not yet been available.
Examination of material from mainland Australia
has demonstrated close similarities to the European
species but with some clear differences. All the
Australian material does not belong to a single species
but represents four different taxa of which three are
new. The differences described below clearly indicate
three distinct taxa, related to previously described
species. The differences are sufficient to require the
creation of three new species.
MATERIAL AND METHODS
Material was mounted in 5% KOH solution
and stained with Congo Red. Specimens are housed
in the J.T. Waterhouse Herbarium, University of New
South Wales (UNSW), except for Western Australian
material, which is in the Western Australian
Herbarium (PERTH). The collections at UNSW all
have extensive field notes and colour photographs
taken under standard conditions.
Spore measurements indicate the range of sizes
found in the various collections. Where spore sizes
are included in brackets, they indicate that the spore
sizes were more than one measuring unit (0.3 um)
beyond the range for all other spores. The value Q
148
represents the mean length:breadth ratio of the spores.
Measurements of Q were averaged for a collection and
where a range is quoted it represents the range across
collections. Measurements of the spores exclude
the apiculus and the ornamentation. Measurements
of cystidia indicate length and maximum width.
Measurements of the basidia exclude the sterigmata.
Colours are usually followed by an annotation
from Maerz and Paul (1950) and have a format such
as 10D3. All colour comparisons were made under
natural light.
The figures show the microscopic features at
standard magnification: spores x2000, cystidia and
basidia x1000. The scale bar represents 10 um at
x2000 magnification.
Key to the SW-Pacific species of Cortinarius
subgenus Cortinarius
1. Average basidiospore length less than 10 um,
cheilocystidia not capitate. . Beate nad
1* Average basidiospore jenothyeh more than 10) um
2. Cheilocystidia 50-140 x 10-25 um,
pleurocystidia scarce, 40-100 x 10-18 um,
lanceolate. UGeIOee A. Seon. C. atroviolaceus
2* Cheilocystidia 30-48 x 4-7 um, pleurocystidia
ABSENT, . PATNI... .OaGee 1.C. jenolanensis
3. Cheilocystidia capitate..........C.austroviolaceus
3* Cheilocystidia not capitate or absent...............4
4. Spores with visible perispore ..............::cceeeceeeee
Pe tah teeth C. subcalyptrosporus
4* Spores without visible perispore....................5
5. Cheilocystidia absent, pleurocystidia rare......
...C. paraviolaceus
SiCheilocystidiaipresentss-..ce-eeee a ee ee 6
6. Spores large, at least up to 12 um ee ee
up to 16 um in length... wee es
6* Spores smaller, at most up 1b 12 um lene 2
slender, Q 1.86, cheilocystidia lageniform, 45-
TOK 12-20 Ree. be eee eC: aly Olagul nus
7. Spores ellipsoid to amygdaliform................. 8
7* Spores broadly ellipsoid to subglobose........... 9
8. Spores elongate Q=1.87, width narrow, 6.3-
7.5 um; cheilocystidia 50-60 x 10-12 um,
pleurocystidia frequent, similar..................
Eee ...2.C. hallowellensis
8* spore ptortee Q- 1. 56, rid broader 7.5-8.5
um; cheilocystidia 35-80 x 15-25 um,
pleurocystidia frequent, similar..... C. violaceus
9. Spores 11-13 x 8-9 um, Q=1.45;
cheilocystidia 55-80 x 14-19 um,
lasenitornas..2) 22 9oe, Aon. ae C. hercynicus
Proc. Linn. Soc. N.S.W., 130, 2009
A.E. WOOD
9* Spores 11-14 x 8.1- 9.3 um, Q=1.40;
cheilocystidia 45 — 120 x 14-17 um, lageniform
PE SOMO are, A LUd, OLA eat 3.C. kioloensis
1. Cortinarius jenolanensis Wood, sp. nov. (Fig.
1: a-d)
Pileo usque ad 4 cm lato, convexo, demum plano,
obscure violaceo, sicco, subtiliter fibrillo-squamoso.
Lamellis obscure violaceis, brunnescentibus. Stipite
ell
5-6 cm longo, 5-8 mm crasso, sicco, appresse
fibrilloso, pallidiori violacea. Sporis 8.4 — 10.2 x 5.7-
6.9 um, Q=1.55, ellipsoideis, subtiliter verrucosis,
cheilocystidiis sparsis, lageniformis 30-48 x 10-
14 wm, absentibus pleurocystidiis, absentibus
pileocystidiis. Hyphis fibuligeris. Habitato in humo
in silvis Eucalyptus mixtis.
Pileus to 4 cm, hemispherical at first, then convex
to flat convex and finally plane, very finely to a little
coarsely radially fibrillose, deep violet, dry, not
i!
Figure 1. Cortinarius jenolanensis (UNSW 88/107) : a. basidiome (x 1); b.spores; c.basidia; d. cheilo-
cystidia; Cortinarius kioloensis (UNSW 83/781) e. basidiome( x 1) f. pileocystidia.
Proc. Linn. Soc. N.S.W., 130, 2009
CORTINARIUS Fr. SUBGENUS CORTINARIUS IN AUSTRALIA
hygrophanous. Lamellae broadly adnate to slightly
decurrent, thin, crowded, with one to two series of
lamellulae, deep violet then deep ferruginous, margin
concolorous. Stipe 50-60 x 5-8 mm, central, firm to
tough, equal to slightly swollen below, sometimes
slightly tapering at the base, upper part cap coloured
or slightly paler, lower part a little paler with base pale
violet, with no obvious basal mycelium, and no clear
zone of velar remains. The only velar remains were a
few scattered appressed fibrils throughout with only
small areas or groups.
Aroma
There is no apparent aroma.
Spores
8.4-10.2 x 5.7-6.9 um, mean 9.44 x 6.09 um,
mean Q = 1.55, oval, suprahilar depression not clearly
present and not clearly smooth, ornamentation low
to very low, a little blunt, not anastomosing. Basidia
25-32 x 11-14 um, clavate, four-spored; clamp
connections present. Cheilocystidia fairly sparse,
variously lageniform (some somewhat irregular)
30-48 x 10-14 um, pleurocystidia absent. Pileal
cuticle a loose layer of narrow hyphae, each 4-7 um
diameter, not encrusted with pigment, mainly radially
arranged and repent, a few a little irregularly loose
and more or less upright with rounded terminal cells
but not specialised as pileocystidia. Below this layer
was a densely packed layer of parallel hyphae, the
layer about 40-50 um thick with individual hyphae
of 4-10 um diameter. Below this layer was a layer
of interwoven hyphae, somewhat compact, of pale
golden hyphae with individual hyphae of 5-8 um
diameter.
Habitat
On soil in eucalypt sclerophyll forest.
Commentary
This species is different from all the species
described by Moser (1987) because of the smooth
pileus, different structure of the cuticle, absence of
pleurocystidia, without amorphous deposits and also
by being of smaller general size and lacking aroma.
It is close to the typical forms of Cortinarius
atroviolaceus but differs in having slightly smaller
spores which are more finely rough and lack a clearly
visible plage, the complete absence of pleurocystidia
and smaller cheilocystidia. It may be that Corner
Collection RSNBB 5258B, noted by Moser(1987),
which has finer ornamentation on the spores and
smaller cheilocystidia, also represents this species.
Cortinarius austroviolaceus is also close, but that
150
species has cheilocystidia that are regularly slightly
capitate and are more variable otherwise, and it also
has a different cuticle with occasional lanceolate
(lageniform) terminal cells. (See Moser 1987, pp
139,140).
Material Examined
NSW : Jenolan Caves, Binda Cabins, Eucalypt
woodland, 30.4.88, A.E.Wood et al. (UNSW 88/107)
Holotype; ACT, Canberra, Tidbinbilla Nature
Reserve, Eucalypt woodland, 16.5.92, A. E. Wood et
al. (UNSW 92/121).
2. Cortinarius kioloensis Wood, sp. nov. (Fig. 1:
e,f; 2: a-c)
Pileo usque ad 6 cm lato, convexo, demum plano,
obscure violaceo, sicco, fibrilloso-squamoso. Lamellis
obscure violaceis, brunnescentibus. Stipite 8-12 cm
longo, 15 mm crasso, basi clavatus usque ad 30 mm
crasso, sicco, appresse fibrilloso, pallidiori violacea.
Sporis 11.1 - 13.5 x 8.1 — 9.3 (10.5) um ellipsoideis.
verrucosis, cheilocystidiis lageniformis, 45-120 x 14
—19 um, pleurocystidiis sparsis, lageniformis 45-113
x 15 —26 um, pileocystidiis cylindricis vel fusiformis
35-60 x 13 —28 um. Hyphis fibuligeris. Habitato in
humo silvis Eucalyptus mixtis.
Pileus to 6 cm diam., rounded convex at first,
then rounded umbonate to convex, finally almost
plane with age, strongly fibrillose to a little tomentose
to finely squamulose, more adpressed with age, deep
violet (48H11—12), becoming blackish with age, dry,
not hygrophanous. Lamellae narrowly to broadly
adnate to slightly sinuate, thin to moderately thick,
somewhat spaced, one or two sets of lamellulae, dark
violet at first, then gradually deep ferruginous, margin
concolorous. Stipe central, firm, solid, bulbous at
base, 8-12 x 1.5 cm, base 3 cm, dry, mostly with clear
fibrillar velar zone and scattered fibrils below, violet
above, somewhat paler than cap, a little paler below
(to 46E6 - 17E4), basal bulb globose, concolorous.
Flesh whitish to pale violet, outer layer of stem dark
violet, deep violet at apex of stipe.
KOH (5%) on cap bright red.
Aroma
Clearly absent even when quite young and fresh;
one collection with slight aroma of wood shavings
(but not camphor wood).
Spores
11.1-13.5 x 8.1-9.3 (10.5) um, mean 12.5
x 8.8 um, Q = 1.37-1.46, grand mean Q = 1.42,
Proc. Linn. Soc. N.S.W., 130, 2009
A.E. WOOD
c
Figure 2. Cortinariu kioloensis (UNSW (83/781) : a. spores; b. basidia; c. cheilocystidia
ovoid to elliptic, suprahilar depression not marked
but present in some cases, but not clearly smooth,
ornamentation moderate, coarse, blunt, with some
slight anastomosing. Basidia 35—50 x 10—12 um, four-
spored; clamp connections present. Cheilocystidia
abundant, ventricose to lageniform, 45-120 x 14-19
uum; pleurocystidia sparse but clearly present, similar
to cheilocystidia, but with some a little fusoid, 45-113
x 15—26 um. Pileal cuticle a layer of loose hyphae
with upturned terminal cells which are somewhat
inflated, swollen or cylindrical, 35-60 x 13-28 um;
subcuticular layer of subcellular cells, 30-40 um
diameter, walls not coloured, below this a narrow
Proc. Linn. Soc. N.S.W., 130, 2009
layer of somewhat inflated, closely packed hyphae,
20-25 um diameter, with coloured contents, below
this the context was of loosely arranged somewhat
inflated hyaline hyphae, 15—25 um diameter.
Habitat
On soil in eucalypt sclerophyll forest.
Commentary
This species is different from the typical forms
of Cortinarius violaceus and C. hercynicus and from
all the other species described by Moser (1986). It
is distinct because of the different habit, absence of
CORTINARIUS Fr. SUBGENUS CORTINARIUS IN AUSTRALIA
aroma, relative scarcity of pleurocystidia, presence
of pileocystidia and spores which are without a well-
differentiated plage and have less well developed wall
ornamentation. There are also some slight differences
in the size and shape of the spores. In this species, the
size and shape are nearer to that found in Cortinarius
hercynicus rather than that found in Cortinarius
violaceus but the shape seems distinctly different
from that of spores of Cortinarius hercynicus in that
the spores are broadly ellipsoid rather than distinctly
amygdaliform. Because of all these features it is
regarded as a distinct taxon and is described as a new
species of Cortinarius near to C. hercynicus.
Material Examined
NSW: Sydney, Scotland Island, Eucalypt
woodland, 22.6.80, S. Lowry, (UNSW 80/268);
Batemans Bay, Kioloa State Forest, Eucalypt
woodland, 19.5.83, A. E. Wood & J. J.Bruhl, (UNSW
83/781) Holotype; Sydney, Royal National Park,
Eucalypt woodland, 5.6.83, F. K. Taeker, (UNSW
83/923);Batemans Bay, Kioloa State Forest, Higgins
Creek, Eucalypt woodland, 15.5.84, A. E. Wood & N.
B. Gartrell, (UNSW 84/495); Sydney, Royal National
Park, Couranga Track, Eucalypt woodland, 28.5.86,
F. K. Taeker, (UNSW 86/254); Sydney, Boronia
Park, Eucalypt woodland, 27.5.90, R. Kearney,
(UNSW 90/197); Hazelbrook, James Park, Eucalypt
woodland, 30.5.92, A.E.Wood et al., (UNSW 92/206);
Springwood, Sassafras Gully, Eucalypt woodland,
16.4.94, A. E. Wood et al.,(UNSW 94/47); Sydney,
Sydney Harbour National Park, Bradleys Head,
7.6.98, B. J.& N. W. Rees, (UNSW 98/25); Sydney,
Lane Cove Bushland Park, Gore Creek, Eucalypt
woodland, 7.6.98, B. J. & N.W. Rees, (UNSW
98/28).
Authentic material from Sweden (Femsjo) was
collected and at first was identified as Cortinarius
violaceus. However later detailed examination clearly
showed that it was a typical example of Cortinarius
hercynicus and the following microscopic details are
added for this collection (as Cortinarius hercynicus
var hercynicus)
Spores
12.6-15.0 x 8.4-9.3 um, mean 13.47 x 8.94 um,
Q=1.51, spores elliptic, only vaguely amygdaliform,
with only some spores showing a slightly flatter
supra-hilar depression, but that mostly not smooth,
ornamentation moderate, a little broad and only
slightly blunt. Cheilocystidia frequent 75-85 x 13-19
uum, narrowly lageniform, pleurocystidia sparse but
clearly present, lageniform, somewhat more variable,
152
50-90 x 12-20 um. Pileal cuticle of closely packed
and interwoven hyphae, layer 100-200um deep,
individual hyphae 5-7 um diameter, without any
terminal cystidia (Fig. 3).
Material Examined :
SWEDEN: Femsjo, woodland, 2.9.79,
M.M.Moser & A.E.Wood, in UNSW(UNSW 79/29).
3. Cortinarius hallowellensis Wood,
sp. nov. (Fig. 4)
Pileo usque ad 6 cm lato, convexo, demum plano,
obscure violaceo, sicco, subtiliter fibrilloso-squamoso.
Lamellis obscure violaceis, brunnescentibus. Stipite
cylindrico vel clavato, 4-7 cm longo, 10-15 mm crasso,
basi leviter, sicco, fibrilloso violacea. Sporis 11.1-
12.0 x 6.3-7.5 um, ovoideo-ellipsoideis, verrucosis,
cheilocystidiis fusiformis vel lageniformis, 50-60 x
9-13 um, pleurocystidis fusiformis, 50-60 x 9-13
um, absentibus pileocystidiis. Hyphis fibuligeris.
Habitato in humo in silvis Eucalyptus mixtis.
Pileus to 3.4-6.0 cm, rounded convex at first,
flattening at maturity, finely radially fibrillose,
very dark violet brown (16F4), not hygrophanous.
Lamellae broadly adnate to adnate, thin, a little
spaced, dark violet (16B5), more rusty with age, with
two series of lamellulae. Stipe cylindrical to clavate,
with a swollen base 3.7-7.0 x 1.0-1.5 cm, solid, dry,
dark violet (16B4) with fine cobweb veil, rapidly
disappearing (after Bougher & Syme 1988, colours
from Kornerup & Wanscher, 1978).
Spores
11.1-12.0 x 6.3-7.5 um, mean 11.49 x 6.81 um,
mean Q = 1.69, oval to elliptic, occasionally vaguely
amygdaliform, with occasionally a slight supra-hilar
depression, but not visibly smooth, ornamentation
moderate, coarse, blunt. Basidia cylindrical to
clavate, 40-55 x 10-12 um, four-spored, clamp
connections present. Cheilocystidia plentiful, narrow
lageniform to fusoid, 50-60 x 9-13 um, pleurocystidia
sparse, but clearly present, similar to cheilocystidia,
but mostly fusiform 50-60 x 10-12 um. Pileal cuticle
with a surface layer 35 -50(80) um deep, a thin layer
of loosely arranged hyphae, individual hyphae 2.5-
5 um diameter, mainly repent, with no erect hyphae
and no differentiated terminal cells, without wall
encrustation, some walls with pale golden walls;
below this a layer of closely packed cylindrical hyphae
of the trama (35-50 x 7-10 um, some a little larger and
a few pseudoparenchymatous cells present).
Proc. Linn. Soc. N.S.W., 130, 2009
A.E. WOOD
Figure 3. Cortinarius herycynicus (UNSW 79/29) : a. spores; b. basidia; c. cheilocystidia; d. pleuro-
cystidia
Commentary
This species is different from the typical
Cortinarius violaceus in that this species has oval to
elliptic spores (Q = 1.69), rather than amygdaliform
spores, the cuticle does not produce pileocystida,
the cheilocystidia are narrower to fusiform and the
general habit is much smaller. Hence it is regarded as
a close, but distinct species.
Material Examined
WA: Denmark, Mount Hallowell Reserve,
Proc. Linn. Soc. N.S.W., 130, 2009
Eucalypt woodland, 22.5.93, K. Syme. (PERTH 0550
6794), Holotype.
Collection PERTH007775665 also seems to be this
species. However it was collected in a Pinus radiata
plantation. It has spores with size 12-13.8 x 6.6-
7.5 um, mean 13.14 x 7.02 um, Q = 1.87, spores
ovoid to elliptic, some vaguely amygdaliform,
supra-hilar depression sometimes slightly present,
but never clearly smooth. Cheilocystidia abundant,
narrow lageniform to narrow fusiform 85-110 x 10-
153
CORTINARIUS Fr. SUBGENUS CORTINARIUS IN AUSTRALIA
int
oD Oo d
Figure 4. Cortinarius hallowellensis (PERTH 0550 6794) : a. spores; b. basidia; c. cheilocystidia;
d. pleurocystidia.
12 um, pleurocystidia abundant, narrow fusiform of
the same dimensions. Pileal cuticle a thin, scarcely
differentiated layer 20-30 wm deep, composed
of narrow hyphae, 2-5 um diameter, the surface
slightly more loosely arranged, but with no special
terminal cells and no upturned cystidia and then the
underlying tissues gradually becoming more densely
packed. This collection has slightly larger spores
and slightly longer cystidia, but does not otherwise
differ from the previous collection. In the absence of
further collections, this is left as another collection of
Cortinarius hallowellensis. This leaves the question
as to whether this form is a local form which has
transferred to the introduced host or whether it was
introduced with the exotic species, and may occur
elsewhere. Much more extensive collecting may
allow this question to be answered.
154
Material Examined
WA:. North of Jarrahdale, Pinus radiata
plantation, 2.6.76, M. Durack.
( PERTH 00775665).
ACKNOWLEDGEMENTS
The advice and encouragement of the late Professor
M.M.Moser is_ gratefully acknowledged. Grateful
appreciation is given to New South Wales National Parks
and Wildlife Service and State Forests of New South Wales
for permission to collect specimens from areas under their
control. Thanks are also due to PERTH herbarium for the
loan of specimens.
Proc. Linn. Soc. N.S.W., 130, 2009
A.E. WOOD
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155
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Late Ordovician Strophomenide and Pentameride Brachiopods
from Central New South Wales
IAN G. PERCIVAL
Geological Survey of New South Wales, Department of Primary Industries, 947-953 Londonderry Road,
Londonderry, NSW 2753, Australia (ian.percival@dpi.nsw.gov.au).
Percival, I.G. 2009. Late Ordovician Strophomenide and Pentameride Brachiopods from central New
South Wales. Proceedings of the Linnean Society of New South Wales 130, 157-178.
Strophomenide and pentameride brachiopods are described from shelfal environments (BA 3) flanking
islands of the Macquarie Arc during the Late Ordovician (latest Sandbian to early Katian stages). Most of
the strophomenoid genera recognized are new, monotypic, and hence endemic, although the occurrence
of a new species of Shlyginia is indicative of affinities with Kazakhstan. Taxa described include the
strophomenid Geniculomena barnesi gen. et sp. nov., the rafinesquinid Testaprica rhodesi gen. et sp. nov.,
glyptomenids Resupinsculpta cuprafodina gen. et sp. nov., Paromalomena zheni sp. nov., and Platymena?
sp., and the plectambonitoid Shlyginia rectangularis sp. nov. Review of the generic assignment of Oepikina?
walliensis Percival, 1991 suggests that this species is better placed in Murinella Cooper, 1956. Relatively
rare pentameride brachiopods are represented by only a few specimens, including an unnamed species of
Parastrophina, and a species tentatively referred to Eoanastrophia.
Manuscript received 1 December 2008, accepted for publication 16 February 2009.
KEYWORDS: brachiopod, Late Ordovician, Macquarie Arc, new genera, pentameride, strophomenide
INTRODUCTION
Late Ordovician strophomenide brachiopods
are well-represented in limestones and sandstones
deposited around volcanic islands forming the
Macquarie Arc in central New South Wales, with most
of the fauna having previously been described over
the past three decades (Percival 1979a, 1979b, 1991;
Percival et al. 2001). For various reasons (including
rarity of specimens, and insufficient knowledge of
morphological features needed to characterize new
species), several additional strophomenide taxa have
remained undocumented. This paper aims to address
this deficiency in order to present a more complete
picture of the fauna to underpin future analyses of
biogeographic relationships. In addition, species
of Late Ordovician strophomenides previously
tentatively ascribed to Oepikina by Percival (1979b)
from the vicinity of Gunningbland, and Percival
(1991) from the Licking Hole Creek area, near
Cliefden Caves (Figure 1), are reassessed in order to
clarify their systematic position.
The opportunity is also taken to describe some rare
examples (represented by just a handful of specimens)
of Late Ordovician pentameride brachiopods. Both
genera recognized are left in open nomenclature as all
specimens are incomplete. However, the presence in
the fauna of two additional camerelloids is significant
and worthy of documentation as only one species of
pentameride brachiopod, Didymelasma inconspicua
Percival, 1991, had previously been described from
contemporaneous rocks of the region.
Except for specimens of Testaprica rhodesi gen.
et sp. nov. and Platymena? sp. which were found in
fine-grained sandstone in the upper Gunningbland
Formation of late Eastonian (Ea3-4) age, the
brachiopods described here are silicified, having
been recovered from residues of limestones dissolved
in dilute hydrochloric acid. These limestones are of
early Eastonian age, equivalent to the latest Sandbian
or earliest Katian of international usage. Details of the
stratigraphic succession and tectonic context within
the Macquarie Arc in central NSW are provided by
Percival and Glen (2007), and only a brief summary
of the age and correlation of these strata (Figure 2) is
given here.
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
BeeeT Gate o
LSI... # BILLABONG
: id CREEK
unningbland ‘ LIMESTONE
e PARKES
e FORBES
e Eugowra
j
| NEW SOUTH WALES $
i
ie
Bathurst
EE ~ oo
N
CANBERRAe
N
Molong |
~REEDY CREEK
/ LIMESTONE
/
Cudal
°
aw
~ BOWAN PARK
CARGO CREEK SHEGREA
LIMESTONE ,
\ Cargo
ORANGE e
| __- REGANS CREEK
"7 LIMESTONE
CANOMODINE — Y ‘
LIMESTONE
e Canowindra
eae
Licking Hole Creek —
CLIEFDEN CAVES
~— LIMESTONE
SUBGROUP
®
Mandurama
® Woodstock
Figure 1. Locality map showing sites in central New South Wales yielding Late Ordovician brachiopods
described in this paper. Outcrop of main Upper Ordovician limestone units shown in black; localities
(L37, L51) in overlying Upper Ordovician clastic-dominated units are shown by spots.
Stratigraphic setting
Cliefden Caves and Licking Hole Creek areas, east
flank of Molong Volcanic Belt
In the Cliefden Caves area of central New South
Wales (Webby and Packham 1982) and the Licking
Hole Creek area adjacent to the west (Percival 1976), a
well preserved Late Ordovician carbonate-dominated
sedimentary succession formed on an eroded volcanic
island setting, represented by the Walli Volcanics.
The Cliefden Caves Limestone Subgroup includes
the Fossil Hill Limestone at the base, succeeded
by the massive Belubula Limestone which is itself
overlain by the Vandon Limestone. Biostratigraphic
evidence from conodonts, trilobites, corals and
stromatoporoids, and brachiopods, demonstrates that
the Fossil Hill Limestone (and equivalents in the
Licking Hole Creek area), and the lower part of the
Belubula Limestone, were deposited in the earliest
Eastonian (Eal); the remainder of the limestone
succession is of Eastonian 2 age, which corresponds
to the basal Katian stage.
158
The strophomenide biofacies characterizes
Benthic Assemblage 3 (BA 3) throughout these
limestone deposits, which is interpreted as occupying
open shelf environments in well-circulated shallow to
moderate water depths (Percival and Webby 1996).
Representative brachiopods of this biofacies have
been largely documented by Percival (1991); further
species described herein include Geniculomena
barnesi, Resupinsculpta cuprafodina, Paromalomena
zheni, Shlyginia rectangularis, and Parastrophina
sp. Additionally Oepikina walliensis Percival, 1991,
described from the basal Belubula Limestone in the
Licking Hole Creek area, is reassessed and assigned
to Murinella.
Regans Creek Limestone, southeast of Cargo, east
flank of Molong Volcanic Belt
The Regans Creek Limestone, mapped by
McLean (1974), is a relatively small exposure of
limestone that is contemporaneous with the Cliefden
Caves Limestone Subgroup.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
BOLINDIAN
Bot Bo2 Bo3 Bo4 Bod
EASTONIAN
wee 1, 2,3
au 1, 2,3
mmm 1, 2,3
mes 1, 2
UPPER ORDOVICIAN
SANDBIAN
Gi2
GISBORNIAN
Gil
Murinella walliensis
Geniculomena bamsei
Resupinsculpta cuprafordina
Paromalomena zheni
Shlyginia rectangulans
Testaprica rhodesi
Platymena? sp.
mu 27,3
Parastrophina sp.
3
Bowan Park
asses le Saeetell 4
ANGULLONG
VOLCANICS
MALACHIS
HILL
t+ FORMATION
MALONGULLI
FORMATION |
GUNNINGBLAND
FORMATION Jee eee
voy te Je
| sieeg a | aeons
| CAVES
LIMESTONE Sleleigaboed
SUBGROUP
BILLABONG
CREEK
LIMESTONE Se SS al
es CARGO WALLI
et VOLCANICS VOLCANICS
=
g
7)
S
& (base early
Wy | (base unknown) (base unknown) Darriwilian)
2009_02_0042
Figure 2. Stratigraphic levels at which Late Ordovician brachiopods described in this paper occur in
central New South Wales. Numerals associated with approximate ranges refer to numbered stratigraph-
ic columns to the right. Note that Parastrophina sp. also occurs in the Checkers Member in the upper
Regans Creek Limestone (not shown on this diagram). HIRN. = Hirnantian stage. :
The Checkers Member in the upper part of the
Regans Creek Limestone yields a silicified fauna
comparable to that in the Trilobite Hill Limestone
Member of the Vandon Limestone at Cliefden Caves,
although diversities are considerably lower. To the
brachiopods described from this level by Percival
(1991) can now be added Parastrophina sp.
Bowan Park area, west flank of Molong Volcanic
Belt
The geology of the Bowan Park area has been
described in detail by Semeniuk (1970, 1973).
Limestones of the Bowan Park Subgroup (including
in ascending order, the Daylesford Limestone,
Quondong Limestone, and Ballingoole Limestone)
overlie the Cargo Volcanics, and are in turn overlain by
Proc. Linn. Soc. N.S.W., 130, 2009
the Malachis Hill Formation (Fig. 2). The succession
at Bowan Park differs from that on the southwestern
MVB (in the Cliefden Caves area) where late
Eastonian (Ea3) age sediments are represented by the
graptolitic Malongulli Formation above the Cliefden
Caves Limestone Subgroup, whereas carbonate
deposition (Ballingoole Limestone) occupied this
interval in the Bowan Park area.
The Quondong Limestone contains abundant
marine invertebrate faunas of the strophomenide
biofacies (Percival 1991), comparable in age and
diversity with those in the Trilobite Hill Member of
the Vandon Limestone (Cliefden Caves Limestone
Subgroup) and like that unit clearly belongs to BA 3
(i.e. shelfal). Additional species described herein from
the Quondong Limestone include Resupinsculpta
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
cuprafodina, Paromalomena zheni, Shlyginia
rectangularis, Parastrophina sp. and Eoanastrophia?
sp.
Gunningbland area, Junee-Narromine Volcanic Belt
The Billabong Creek Limestone was shown
by Pickett and Percival (2001) to extend from
southeast of Gunningbland in a broad arcuate band
trending northwestwards to north of the Parkes-
Broken Hill railway, then northeast to exposures
on “Kirkup” property (Figure 1). Conodonts from
the “Kirkup” section, of early Darriwilian (Da2)
age (Zhen and Pickett 2008), are the oldest dated
fossils in the Billabong Creek Limestone. Younger
conodont and coral assemblages from the type
section of the formation on “Nelungaloo” property,
southeast of Gunningbland, range in age through the
late Darriwilian, Gisbornian and earliest Eastonian
(Pickett and Percival 2001). Outcrops in and adjacent
to Billabong Creek at the southern extremity of the
limestone belt are rich in silicified fossils, particularly
brachiopods (including Geniculomena barnesi,
Resupinsculpta cuprafodina, Paromalomena zheni
and Shlyginia rectangularis, described herein, and
a diverse fauna documented by Percival 1991) and
trilobites (Webby 1973, 1974), of Eastonian 2 age
(Pickett and Percival 2001). These upper beds of
the Billabong Creek Limestone correlate with the
Quondong Limestone at Bowan Park, and the Trilobite
Hill Limestone Member of the Vandon Limestone in
the Cliefden Caves Limestone Subgroup (Figure 2).
The Billabong Creek Limestone is apparently
conformably overlain by the Gunningbland
Formation, although the actual boundary is unexposed.
The outcrop belt of the Gunningbland Formation
consistently lies immediately west of the arcuate
trend of the Billabong Creek Limestone exposures
(Pickett and Percival 2001). Shallow excavations and
exposures in ploughed fields on “Currajong Park”,
“Sunnyside” and “New Durran” properties in the
Gunningbland district reveal that the Gunningbland
Formation predominantly consists of siltstone, shale,
and fine- to medium-grained sandstone, together with
minor fossiliferous limestones.
Most of the Gunningbland Formation is of late
Eastonian (Ea3) age, determined from graptolites in
siltstones, and conodonts including Taogupognathus
tumidus in limestone lenses. The limestones
also contain a coral-stromatoporoid assemblage
corresponding to the contemporaneous Fauna III
(McLean and Webby 1976, Webby and Morris 1976).
Two brachiopod faunas, elements of which were
described by Percival (1978, 1979a, 1979b), are
recognised. Brachiopod Fauna C, of Ea3 age, is present
160
in the lower part of the formation on “New Durran”
property. The presumed latest Eastonian age of Fauna
D (Percival 1992), occurring in strata on “Currajong
Park” property, was confirmed by the presence
of graptolites of Ea4 age in the uppermost beds of
this section (Pickett and Percival 2001). A diverse
trilobite fauna has recently been described from this
upper part of the unit (Edgecombe and Webby 2006,
2007), associated with the brachiopods Testaprica
rhodesi gen. et sp. nov. and Platymena? sp. which are
documented herein. This completes description of the
brachiopod fauna collected from the Gunningbland
Formation over more than three decades; two other
genera (Christiania sp., Ptychopleurella? sp.) are
represented in the upper part of this unit by single
specimens of ventral valves, which do not warrant
description until further material is forthcoming.
Systematic palaeontology
Type material (designated MMF), comprising
specimens described and illustrated or listed herein,
is curated in the palaeontological collections of the
Geological Survey of New South Wales held at
Londonderry in western Sydney. Some specimens
labeled SUP, including material of Murinella
walliensis and an external mould of the ventral valve
of Testaprica rhodesi, were transferred from the
Geology Department of the University of Sydney
to the Australian Museum, Sydney in the mid-
1980s (these are awaiting renumbering). For brevity,
authorship of taxonomic hierarchy above genus level
is not cited in the References; these bibliographic
sources are listed in the revised (2 edition) Treatise
of Invertebrate Paleontology, Part H: Brachiopoda
Volume 3 (Williams et al. 2000).
Phylum Brachiopoda Duméril, 1806
Subphylum Rhynchonelliformea Williams,
Carlson, Brunton, Holmer and Popoy, 1996
Class Strophomenata Williams, Carlson,
Brunton, Holmer and Popov, 1996
Order Strophomenida Opik, 1934
Superfamily Strophomenoidea King, 1846
Family Strophomenidae King, 1846
Subfamily Furcitellinae Williams, 1965
Geniculomena gen. nov.
Type species (by monotypy): Geniculomena barnesi
gen. et sp. nov.
Diagnosis
Dorsally geniculate planoconvex to weakly
concavoconvex furcitellin with unequally
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
parvicostellate ornament lacking rugae; teeth and Diagnosis
sockets without crenulations; dorsal myophragm As for genus.
absent; septa associated with dorsal muscle field Etymology
are less strongly developed than single continuous Genus name in reference to geniculate dorsal
median ridge. valve profile and broadly crescent-like shell outline;
species name honours David Barnes, photographer
Geniculomena barnesi gen. et sp. nov. in the NSW Department of Primary Industries, in
Fig. 3 A-N appreciation of the assistance he has provided to me
Figure 3. Geniculomena barnesi gen. et sp. nov. A — B: interior and exterior of dorsal valve, holotype
MMF 44915. C — E: interior, exterior and lateral profile (dorsal side uppermost) of dorsal valve, MMF
44916. F—H: interior, exterior and anterior profile (dorsal side uppermost) of dorsal valve, MMF 44919.
I — J: interior and exterior (bearing heliolitid coral) of dorsal valve, MMF 44917. K — L, O — Q: one
incomplete individual shell, which disarticulated during acid dissolution of limestone matrix; K — L: ex-
terior and lateral profile (dorsal side uppermost) of dorsal valve, MMF 44918a; O — Q: exterior, interior
and lateral profile (ventral side uppermost) of ventral valve, MMF 44918b. M: interior of dorsal valve,
MMF 44920. N: interior of dorsal valve, MMF 44921; note distortion on anterolateral margin, probably
indicating repaired injury. Scale bar below C represents one cm. A— E, I— L, O— Q from L24, Trilobite
Hill Limestone Member of Vandon Limestone, upper Cliefden Caves Limestone Subgroup at Licking
Hole Creek near Walli; F — H from L135 (east of Copper Mine Creek, near Cliefden Caves) in Trilobite
Hill Limestone Member of Vandon Limestone, upper Cliefden Caves Limestone Subgroup; M — N from
L143, upper Billabong Creek Limestone at Billabong Creek road crossing south of Gunningbland.
Proc. Linn. Soc. N.S.W., 130, 2009 161
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
over the past decade in preparing many illustrations
of fossils for publication.
Material
Five dorsal valves, mostly entire, and one partial
ventral valve with corresponding partial dorsal valve
(disarticulated), all material silicified. Holotype is
dorsal valve MMF 44915; paratypes include dorsal
valves MMF 44916, MMF 44917, MMF 44919,
MME 44920 and 44921, and ventral valve MMF
44918a and corresponding dorsal valve 4491 8b.
Localities
Type locality is L24 (Licking Hole Creek area),
in Trilobite Hill Limestone Member of Vandon
Limestone, upper Cliefden Caves Limestone
Subgroup; also found in same stratigraphic unit at
L135 (east of Copper Mine Creek, near Cliefden
Caves); also occurs at L143 in upper Billabong Creek
Limestone, at Billabong Creek road crossing south
of Gunningbland [full details of these localities are
given by Percival 1991].
Description
Shell planoconvex to very weakly concavoconvex
(rarely ventribiconvex, e.g. Fig. 3G), becoming
dorsally geniculate when fully grown; transversely
subquadrate with maximum width either at, or
immediately anterior to, hingeline; lateral and anterior
margins broadly curved. Shell of moderate size,
ranging in length from 12 to 16 mm, and in width from
23 to 29 mm in largest specimens; length to width
ratio 0.55 -0.80. Ornament unequally parvicostellate,
with every fourth or fifth rib accentuated; rugae
lacking; exterior of the sole ventral valve assigned to
this species is almost entirely devoid of ornament, but
this may have been eroded prior to fossilization.
Ventral interior (described from an incomplete
valve) shows robust oblique teeth supported by low
plates for approximately three-quarters length; dental
plates extend anteriorly to bound triangular diductor
scars flanking (but not enclosing) narrower median
pair of adductor scars separated by low median ridge
not extending forward of muscle field which occupies
three-eighths valve length. Mantle canals prominent,
of lemniscate type with anteriorly divergent vascula
media not enclosing vascula genitalia. A distinct but
low subperipheral rim defines a dorsally-deflected
marginal band approximately one-seventh of valve
length extending around entire lateral and anterior
valve margin. Details of interarea and delthyrium not
known.
Dorsal interior with Type A strophomenoidean
cardinalia consisting of twin cardinal process lobes
162
extending just posterior to hingeline and convergent
above a hollow, with narrow, widely divergent socket
ridges recurved posterolaterally at extremities;
sockets short but deep; no crenulations visible on
socket ridges. Notothyrial platform poorly developed,
lacking myophragm; low median septum extends
from immediately in front of cardinal process lobes
to terminate at about half valve length, separating
moderately conspicuous pair of adductor scars which
are bounded by weaker side septa; short transmuscle
septa barely visible or lacking. Mantle canals
apparently lemniscate, poorly expressed, except for
vascula genitalia in largest specimen. A variably
defined subperipheral ridge is sometimes developed
slightly posterior to dorsally-directed geniculation of
marginal band.
Dimensions
Holotype MMF 44915: length 12.0 mm, width
19.0 mm; paratypes MMF 44916: length 13.1 mm,
estimated width 23 mm; MMF 44917: length 15.5
mm, width of specimen (incomplete) 18.8 mm;
MME 44919: length 16.0 mm, width 23.5 mm; MMF
44920: length 13.5 mm, estimated width 22.5 mm;
MMF 44918a (vv): length 15.9 mm, estimated width
29 mm.
Discussion
Geniculomena is assigned to the subfamily
Furcitellinae, rather than the Strophomeninae, due
to the presence of a moderately well-defined dorsal
muscle field in some specimens, although muscle
bounding ridges, side septa and transmuscle septa
are somewhat variably developed and may be barely
discernible in other examples depending on degree
of silicification. Dorsally geniculate genera similar
to Geniculomena are more typical of furcitellins
rather than strophomenins. Dactylogonia Ulrich and
Cooper, 1936 (and its synonym Cyphomena Cooper,
1956) appears to closely resemble Geniculomena
in general morphology, but Dactylogonia is readily
distinguished by its much stronger development of
transmuscle and side septa in the dorsal valve. The
new genus lacks the characteristic rugate ornament
of Bellimurina Cooper, 1956, and differs internally in
absence of a forked anterior termination to the dorsal
median ridge.
Although Geniculina R6émusoks, 1993, from
the latest Ordovician (Hirnantian) of the Baltic region,
is broadly similar to Geniculomena, the new genus
apparently lacks the prominent posterolateral oblique
rugae developed on the ventral valve of Geniculina.
Nor have crenulations been observed on the teeth and
socket ridges of Geniculomena, whereas these are
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
characteristic of at least four species of Geniculina
(e.g., ROOmusoks 2004, pl. IX fig. 12, pl. XI fig.
6). The median septum in Geniculomena is a single
ridge that extends from the cardinal process and is
rather more prominent than the side septa, unlike
the arrangement in Geniculina that has strong side
septa and a stout myophragm which bifurcates at its
anterior extremity.
The multicostellate ornament of Maakina
Andreeva, 1961 (in Nikiforova and Andreeva 1961),
from the early Katian of the Siberian Platform, is
quite different from that of Geniculomena. Internally,
the absence of a dorsal median septum and presence
of crenulations on the socket ridges in Maakina are
additional features clearly distinguishing these two
genera.
Distribution
Early Eastonian (Ea2), equivalent to basal
Katian; presently monotypic and known only from
limestones of the Macquarie Arc in central NSW.
Murinella Cooper, 1956
Type species: Murinella partita Cooper, 1956
Murinella walliensis (Percival, 1991)
Fig. 4 A-G
Synonymy
Oepikina? walliensis Percival, 1991: p.147, fig.
14.20-28.
Discussion
Two species with Oepikina-like morphology have
previously been described from the Late Ordovician
of central NSW. One form from the Gunningbland
Formation was tentatively referred to Oepikina? sp.
by Percival (1979b), and a new species Oepikina?
walliensis was described by Percival (1991) from
the Licking Hole Creek area, occurring in strata
equivalent to the basal Belubula Limestone. In their
revision of the superfamily Strophomenoidea, Rong
and Cocks (1994, p.694) noted that the cardinalia
of O? walliensis was “of the Strophomena group”,
presumably implying that in their view the species
was a strophomenin rather than a furcitellin. Zhan
et al. (2008) observed that these two subfamilies
are difficult to separate using the revised Treatise
classification (Cocks and Rong 2000). Rong and Cocks
(1994, text-fig. 3) also presented a well-illustrated
comparison between the dorsal cardinalia of the type
species of Strophomena and Murinella. Although
a reclassification of O? walliensis on the basis of
cardinalia alone might therefore be superfluous, the
comments by Rong and Cocks (1994) have prompted
a reassessment of other possible generic affinities of
Figure 4. Murinella walliensis (Percival, 1991). A— E: Holotype SUP 68516, exterior of conjoined valves,
dorsal, ventral, posterior profile, anterior profile and lateral profile respectively. F: fragment of dorsal
valve interior showing cardinalia, SUP 68523. G: interior of ventral valve, SUP 68518. Both scale bars
represent 1 cm (that beneath F pertains only to this specimen; the shorter scale bar applies to specimens
A-E and G). All specimens from basal Belubula Limestone at Licking Hole Creek, near Walli.
Proc. Linn. Soc. N.S.W., 130, 2009
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
this species.
The holotype of O? walliensis is here refigured,
together with a paratype fragment showing the
cardinalia and a ventral valve interior. Reasons given
by Percival (1991) for provisionally assigning this
species to Oepikina include poorly developed septa
in the dorsal valve, and presence of a relatively small
ventral muscle field enclosed by low bounding ridges.
Both these features are atypical of Oepikina, whereas
they are characteristic of the similar genus Murinella
Cooper, 1956. Although a distinguishing feature of
the type species of Murinella, M. partita Cooper,
1956, is the extension of the median septum anterior
to the ventral muscle field, not all species show this
(e.g. M. muralis Cooper, 1956 and M. semireducta
Cooper, 1956). In retrospect, O? walliensis accords
best with Murinella, and it is here designated M.
walliensis (Percival, 1991). Other features supporting
this reassignment include the relatively large
pseudodeltidium and prominent subperipheral rim
in the dorsal valve of M. walliensis. Furthermore,
the cardinalia definitely conform to the Murinella
model.
A species of Murinella has also been described
from the lower limestone member of the Benjamin
Limestone in Tasmania by Laurie (1991). That
species, M. magna, is distinguished by its much larger
dimensions, and in having a median septum extending
forward of the ventral muscle field, compared to M.
walliensis.
Oepikina? sp from Gunningbland is known only
from one specimen (Percival 1979b, fig. 1.12), which
clearly shows the presence of Type A cardinalia (sensu
Rong and Cocks 1994). In all other features this
dorsal valve is definitely Oepikina-like, with strong
side septa, but the absence of a corresponding ventral
valve continues to prevent a confident assignment
to that genus. The external mould supposedly of a
dorsal valve (SUP 62569), mentioned but not figured
by Percival (1979b, p.183), is now considered to be a
ventral valve of Testaprica rhodesi (see below) rather
than being related to Oepikina.
Family Rafinesquinidae Schuchert, 1893
Subfamily Rafinesquininae Schuchert, 1893
Testaprica gen. nov.
Type species (by monotypy): Zestaprica rhodesi gen.
et sp. nov.
Diagnosis
Convexo-concave to convexo-planar
rafinesquinin similar to Rhipidomena but with
164
prominent subparallel side septa in dorsal valve; other
septa and median ridge subdued or lacking.
Testaprica rhodesi gen. et sp. nov.
Fig. 5 A-H
Diagnosis
As for genus.
Etymology
Genus name derived from testa (Latin): shell,
and apricum (Latin): a sunny spot, in reference to the
occurrence of this brachiopod adjacent to “Sunnyside”
property; species named in honour of Julie and
John Rhodes, former owners of “Sunnyside” and
“Currajong Park” properties at Gunningbland, who
kindly provided access to collect on their land, and
who also recognised and donated several important
brachiopods and trilobites for scientific description.
Material
Holotype: MMF 36806a and b, dorsal valve
internal mould and external mould of corresponding
ventral valve. Paratypes: MMF 36798a and b, dorsal
valve internal and external moulds; MMF 36801 and
MME 36805, both external moulds of dorsal valves;
MMF 36813, dorsal valve internal mould; SUP 62569
ventral valve external mould.
Localities
All specimens from upper Gunningbland
Formation on “Currajong Park”, Gunningbland at
locality L51 [see Percival 1979a for full details] with
exception of MMF 36813, collected from locality
L48 situated in immediately underlying beds in the
same formation on this property.
Description
Large convexo-concave to convexo-planar shells
up to 40 mm wide and 30 mm long, with maximum
width attained at or immediately anterior to hingeline;
anterolateral and anterior margins very broadly
rounded. Length to width ratio varies between two-
thirds and almost three-quarters. Ornament finely
and evenly multicostellate, lacking rugae; costellae
slightly curved on lateral flanks; occasional concentric
growth discontinuities may be present, but concentric
filae lacking.
Ventral valve weakly concave, becoming almost
planar anteriorly; interarea low, catacline to weakly
apsacline, with small pseudodeltidium. Details of
interior unknown.
Dorsal valve strongly convex; interarea very low
with delicate chilidial plates (poorly preserved on
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Figure 5. Testaprica rhodesi gen. et sp. nov. All specimens from upper beds of the Gunningbland For-
mation on “Currajong Park”, Gunningbland. A — D: Holotype, MMF 36806a and b; A: exterior mould
of ventral valve (on left, 36806a) and interior mould of corresponding dorsal valve (on right, 36806b);
B: latex replica taken from this specimen; C: enlargement of posterior region of latex replica of dorsal
valve; D: latex replica of exterior of ventral valve, tilted to better show ornament and interarea. E: inte-
rior mould of dorsal valve, MMF 36798a. F: latex replica of dorsal valve, MMF 36813. G: latex replica of
exterior of dorsal valve, MMF 36805. H: latex replica of exterior of dorsal valve, MMF 36801. Both scale
bars represent 1 cm (that below C pertains only to this enlargement).
available specimens). Cardinalia consisting of small transmuscle septa barely visible; muscle bounding
cardinal process with pair of discrete peg-like lobes ridges not present and muscle field not impressed.
above low notothyrial platform, with very short, Mantle canals not discernible.
straight socket ridges extending obliquely; median
ridge either very short or not developed; prominent Dimensions
subparallel pair of side septa, low and thin, extend MME 36806a, b (holotype): DV internal mould
to between one quarter and one third valve length; and VV external mould L= 26.3 mm, hinge W= 39.3
mm;
Proc. Linn. Soc. N.S.W., 130, 2009 165
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
MMF 36798a, b: DV internal and external
moulds L= 28.4 mm, W= 39.3 mm;
MME 36801: DV external mould L= 22.4 mm,
spec W= 25.3 mm, W= 29.6 mm;
MME 36805: DV external mould L= 16.0 mm,
W= 22.0 mm;
MME 36813: DV internal mould L= 17.6 mm;
SUP 62569: VV external mould L= 13.5 mm,
W= 17.5 mm.
Discussion
This monotypic genus has cardinalia of Type B
(sensu Rong and Cocks 1994), with small discrete
cardinal process lobes that are not continuous with a
median ridge, and which are also definitely disjunct
from the socket ridges (the latter being straight
and oblique, rather than recurved laterally towards
the hingeline as in strophomenids). Clearly then,
its affinities lie with the rafinesquinids. The only
previously described rafinesquinin brachiopod with
a convexo-concave valve profile is Rhipidomena,
which is of generally comparable size. However,
dorsal valves of the 5-6 species of this genus known
from North America (Cooper 1956), are never
quite as convex as is Testaprica, and the latter
is not resupinate as is commonly the case with
Rhipidomena. In possessing prominent side septa T-
rhodesi differs from all North American Rhipidomena
species, and is further distinguished by its relatively
poorly developed median ridge and transmuscle septa
(although there is some variation in the strength of
these features). These distinctions in total appear to
be of generic significance, so that despite the absence
of ventral interiors the establishment of a new genus
is warranted.
Equally prominent side septa are also
characteristic of Lateriseptomena Zhan, Jin, Rong,
Chen and Yu, 2008, known from two species of late
Katian age from Zhejiang Province, south-east China.
However, Lateriseptomena has Type C (glyptomenid)
cardinalia, and furthermore has a _ planoconvex
to biconvex profile, so is apparently not closely
related to Testaprica. The concavo-convex profile of
Dirafinesquina Cocks and Zhan, 1998, from Upper
Naungkangyi Group equivalent strata (probable
Katian age) in the Southern Shan States of Burma,
readily distinguishes this genus from Testaprica; the
few known dorsal interiors of Dirafinesquina also
lack the characteristic side septa of the new genus.
Distribution
Presently known only from the Gunningbland
Formation (upper part) in vicinity of Gunningbland
village, between Parkes and Bogan Gate, central west
166
NSW; late Eastonian (Ea3-4) i.e. Katian.
Family Glyptomenidae Williams, 1965
Subfamily Glyptomeninae Williams, 1965
Resupinsculpta gen. nov.
Type species (by monotypy): Resupinsculpta
cuprafodina gen. et sp. nov.
Diagnosis
Resupinate glyptomenin displaying weak
rugation on exterior of both valves; teeth and socket
ridges occasionally crenulate.
Resupinsculpta cuprafodina gen. et sp. nov.
Fig. 6 A-P
Diagnosis
As for genus.
Etymology
Genus name in reference to resupinate profile and
finely engraved appearance of ornament (resupinus:
L bent back; insculptus: L engraved); species name
in reference to Copper Mine Creek, the type locality
(cuprum: L copper; fodina: L mine or pit).
Material
Holotype MMF 44923 (conjoined valves);
paratypes include MMF 44924 (ventral valve), MMF
44925 (dorsal valve), MMF 44926 (ventral valve),
MMF 44927 (dorsal valve), MMF 44928 (ventral
valve), and MMF 44929 (conjoined valves). All
specimens are silicified.
Localities
Type locality L135 (east of Copper Mine
Creek, near Cliefden Caves), in Trilobite Hill
Limestone Member of Vandon Limestone, upper
Cliefden Caves Limestone Subgroup; also found at
L138 (“Quondong”, Bowan Park, east of Cudal) in
Quondong Limestone, Bowan Park Subgroup; and at
L144 in upper Billabong Creek Limestone beside the
road crossing Billabong Creek, south of Gunningbland
[full details of these localities are given by Percival
NSO:
Description
Shell relatively small, length up to 12 mm and
width to approximately 18 mm; outline subquadrate
initially, becoming transverse and slightly auriculate
when fully grown with maximum width at hinge line;
length two-thirds width in these largest specimens.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Figure 6. Resupinsculpta cuprafodina gen. et sp. nov. A— D: Holotype conjoined valves, MMF 44923;
A: exterior of ventral valve; B: exterior of dorsal valve; C: lateral profile (dorsal valve uppermost); D:
posterior profile (dorsal valve uppermost). E — F: exterior and interior of ventral valve, MMF 44924.
G — I: exterior and interior of ventral valve, and enlargement of delthyrium to show crenulated teeth,
MMF 44926. J — K: exterior and interior of dorsal valve, MMF 44925. L — M: interior and exterior of
dorsal valve, MMF 44927. N: exterior of ventral valve, MMF 44928. O — P: conjoined valves, ventral and
dorsal exteriors respectively, MMF 44929. Scale bar represents 1 cm for whole figure (except I, which is a
five-times enlargement of H). A— F, J — K from L135 (east of Copper Mine Creek, near Cliefden Caves):
in Trilobite Hill Limestone Member of Vandon Limestone, upper Cliefden Caves Limestone Subgroup;
G —-I, N — P from L138 (“Quondong”, Bowan Park, east of Cudal) Quondong Limestone, Bowan Park
Subgroup.
Ventral valve with sharply pointed beak; profile
initially weakly convex, becoming resupinate in
largest specimens; dorsal valve planar posteriorly,
gently to moderately convex anteriorly in adults;
whole shell very compressed dorsoventrally.
Ornament unequally parvicostellate, commonly with
3-4 finer costellae between accentuated ribs, with
indistinct rugae developed posteriorly.
Ventral interarea low, apsacline, with wide
delthyrium covered apically by pseudodeltidium.
Delicate teeth, crenulated in one specimen (Fig.
Proc. Linn. Soc. N.S.W., 130, 2009
61), supported by thin subparallel dental plates that
terminate immediately in front of teeth. Muscle field
indistinct, apparently very short, not enclosed by
ridges. A weak subperipheral rim is present in one
specimen. Mantle canals not visible.
Dorsal interarea very low, orthocline to weakly
anacline; notothyrium entirely occupied by cardinal
process lobes; chilidial plates either lacking or
extremely weakly developed. Cardinalia consist of
small paired cardinal process lobes fused to long,
straight, widely divergent socket ridges (which are
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
finely crenulated in at least one specimen, Fig. 6L)
with slightly curved terminations; cardinal process
lobes extend very slightly posteriorly of hingeline
and anteriorly overhang a concavity in place of
notothyrial platform; median ridge short, very low;
side and transmuscle septa absent. Muscle field and
mantle canals not visible.
Dimensions
Nearly all specimens are incomplete; a juvenile
conjoined shell MMF 44929 is 6.2 mm long and 7.7
mm wide; the largest shell (holotype, MMF 44923) is
11.6 mm long and 18.5 mm wide.
Discussion
The new species presents a conundrum as regards
its generic affinities. It has Type C (glyptomenin)
cardinalia, and conforms in almost all respects with
the characteristics of Glyptomena, except for the
resupinate profile of larger shells. Smaller shells are
planoconvex and thus more similar to the typical
concavoconvex profile of Glyptomena. As shell profile
is often used to distinguish genera in strophomenides,
it seems reasonable to establish a new genus within
the glyptomenines based on the resupinate character.
Furthermore, crenulated teeth and socket ridges as
seen in Resupinsculpta cuprafodina are apparently
rare in glyptomenines; Rong and Cocks (1994) only
mentioned their occurrence in Mjoesina, which
was doubtfully assigned to the family (Cocks and
Rong 2000), but is now regarded more likely to be
a rafinesquinid (Cocks 2005). The indistinct rugae
present in the posterior region of the exterior of both
valves of R. cuprafodina are lacking in species of
Glyptomena, but the distinctively dorsally geniculate
Glyptomenoides Popov and Cocks, 2006 (which
is otherwise generally similar to Glyptomena) also
displays irregular rugae. Most comparable of other
strophomenids is possibly Longvillia Bancroft, 1933,
which also is resupinate; however, Longvillia has
Type A cardinalia and is therefore not closely related
to the new genus.
Distribution
Only known from limestones of early Eastonian
(Ea2) age, equivalent to the earliest Katian Stage, in
the Macquarie Arc, central NSW.
Paromalomena Rong, 1984
Type species: Platymena polonica Temple, 1965
Paromalomena zheni sp. nov.
Fig. 7 A-V
168
Diagnosis
A species of Paromalomena distinguished by its
prominent pseudodeltidium with a minute foramen at
the apex, and lacking conspicuous external rugae.
Etymology
This species is named in honour of my colleague
Dr Yong-Yi Zhen, in recognition of his extensive
palaeontological studies in the Ordovician of both
Australia and China.
Material
Holotype is MMF 44932 (ventral valve);
paratypes include MMF 44930 (dorsal valve), MMF
44931 (ventral valve), MMF 44933 (ventral valve),
MME 44934 (conjoined valves), MMF 44935 (dorsal
valve), MMF 44936 (ventral valve), MMF 44937
(conjoined valves), MMF 44938 (dorsal valve),
MMEF 44939 (dorsal valve), MMF 44940 (dorsal
valve), MMF 44941 (dorsal valve), MMF 44942
(dorsal valve), MMF 44943 (ventral valve), MMF
44944 (ventral valve), MMF 44945 (ventral valve),
and MMF 44946 (conjoined valves). All specimens
are silicified.
Localities
Type locality is L138 (“Quondong”, Bowan Park,
east of Cudal) in Quondong Limestone, Bowan Park
Subgroup; also occurs at L24 (Licking Hole Creek
area, Walli) in Trilobite Hill Limestone Member of
Vandon Limestone, upper Cliefden Caves Limestone
Subgroup; and at localities L143 and L144 in upper
Billabong Creek Limestone, in vicinity of Billabong
Creek road crossing, south of Gunningbland [full
details of these localities are given by Percival
1991].
Description
Shells generally small and thin, not exceeding 7.5
mm in length and 9.2 mm in width, with subquadrate
to subrectangular outline; hingeline straight and
wide, in all but one specimen just slightly narrower
than maximum valve width which is approximately
coincident with midlength, anterior margin broadly
rounded; length:width ratio ranges from 0.65 to
0.88, with average of 0.77 for 18 specimens. Profile
generally planoconvex, to weakly concavoconvex with
tendency to geniculation dorsally in largest specimens;
a subtle sulcus may develop in anteromedian sector of
dorsal valve, with corresponding weak fold in ventral
valve. Ornament finely and equally parvicostellate,
lacking rugae; occasional concentric growth
discontinuities may be present. Ventral interarea
apsacline, with relatively wide delthyrium at least half
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Figure 7. Paromalomena zheni sp. nov. A — B: exterior and interior of ventral valve, MMF 44931. C: inte-
rior of dorsal valve, MMF 44935. D: exterior of ventral valve, MMF 44936. E — F: exterior and interior of
dorsal valve, MMF 44930. G: interior of ventral valve, holotype MMF 44932. H: interior of ventral valve,
MMEF 44933. I — J: conjoined valves, ventral and dorsal exteriors respectively, MMF 44934. K: dorsal
exterior of conjoined valves, MMF 44937. L: interior of juvenile dorsal valve, MMF 44938. M: interior
of juvenile ventral valve, MMF 44944. N — O: conjoined valves, ventral and dorsal exteriors respectively,
MME 44946. P—Q: exterior and interior of dorsal valve, MMF 44939. R: interior of ventral valve, MMF
44945. S: exterior of dorsal valve, MMF 44940. T: interior of dorsal valve, MMF 44941. U: interior of
dorsal valve, MMF 44942. V: exterior of ventral valve, MMF 44943. Scale bar represents 1 cm. A — L
from L138 (““Quondong”, Bowan Park, east of Cudal) Quondong Limestone, Bowan Park Subgroup; M,
Q-V from L143, upper Billabong Creek Limestone at Billabong Creek road crossing south of Gunning-
bland; N — O from L24, Trilobite Hill Limestone Member of Vandon Limestone, upper Cliefden Caves
Limestone Subgroup at Licking Hole Creek near Walli.
to three-quarters covered by prominent high convex Ventral interior: Pedicle foramen about pin-
pseudodeltidium; a minute pedicle foramen is present hole size, encased in callus at extreme posterior of
at apex of pseudodeltidium. Dorsal interarea barely _delthyrial cavity. Small teeth supported by receding
evident, considerably lower than that of ventral valve; dental plates, below which extend anteriorly
chilidial plates (if present) extremely delicate. divergent, subparallel or slightly convergent lateral
Proc. Linn. Soc. N.S.W., 130, 2009
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
muscle bounding ridges that rapidly decline in
height and do not enclose muscle field anteriorly;
diductors surround adductors that are embedded in
shallow subcircular pit on low median ridge. Muscle
field occupies approximately one-third valve length
and less than one-quarter width. Mantle canals not
observed.
Dorsal interior: Cardinalia of glyptomenin type
(Type C), with very delicate cardinal process lobes
joined to fine, short socket ridges that diverge and
curve to extend subparallel to hingeline; notothyrial
platform absent; low, broad median ridge is barely
developed in some larger specimens, otherwise
lacking; side and transmuscle septa never developed;
muscle scars not clearly defined. Mantle canals not
discernible, due to thinness of shell material that
reflects external costellae.
Dimensions
Valve length ranges from 3.2 mm to 7.5 mm,
and valve width ranges from 4.3 mm to 9.2 mm
(measurements from 18 individuals; no significant
difference between ventral and dorsal valves).
Holotype (ventral valve MMF 449372) is 7.5 mm long
and 9.2 mm wide; majority of specimens cluster in
the range of 5.0-6.5 mm long, and 5.5-8.5 mm wide.
Discussion
This new species shares many morphological
characteristics with the cosmopolitan Late Ordovician
(late Katian — Hirnantian) genus Paromalomena
including shell profile and ornament, development
of fold and sulcus anteriorly, and in most internal
details. It differs from described species mainly in
having a conspicuous pseudodeltidium, and in lacking
a large chilidium and external rugae. Paromalomena
typically occurs in deepwater settings (BA 4-6) in
distinctive faunal associations such as the Foliomena
fauna (e.g. Neuman 1994) and the younger Hirnantia
fauna (e.g. Temple 1965). Like these species, P. zheni
is quite thin-shelled, but unlike them it occurs in
considerably shallower environments (BA 3) and is
somewhat older (earliest Katian).
Unlike species of Glyptomena, the new
species has a furcitellin-like ornament (i.e. equally
parvicostellate), and is generally planoconvex rather
than concavo-convex, except in largest specimens. P.
zheni 1s readily distinguished from Resupinsculpta
cuprafodina, the other glyptomenin with which it is
associated in the same strata in central NSW, by the
latter’s resupinate profile, unequally parvicostellate
ornament and presence of rugae.
Glyptomenoides species differ in having an
unequally parvicostellate ornament with rugae
developed, and furthermore are quite distinct internally
from P. zheni which lacks a stout myophragm and
transmuscle septa.
Distribution
Limestones of early Eastonian (Ea2) age,
equivalent to the earliest Katian Stage, in the
Macquarie Arc, central NSW.
Platymena Cooper, 1956
Type species: Platymena plana Cooper, 1956
Platymena? sp.
Fig. 8 A-D
Material
MMF 36804, external mould of ventral valve;
MMEF 36810, internal mould of dorsal valve; MMF
44968, internal mould of dorsal valve (not figured).
Figure 8. Platymena? sp. A — B: latex replica of ventral valve exterior and interarea of conjoined valves,
MMF 36804. C — D: latex replica and corresponding internal mould of dorsal valve, MMF 36810. All
specimens from upper beds of the Gunningbland Formation on “Currajong Park”, Gunningbland. Scale
bar represents 1 cm.
170
Proc. Linn. Soc. N.S.W., 130, 2009
1.G. PERCIVAL
Locality
All three known specimens from sandstones in
upper Gunningbland Formation on “Currajong Park’,
Gunningbland at locality L51 [see Percival 1979a for
full details].
Description
Transverse auriculate shell with maximum width
at hingeline; lateral and anterior margins broadly
rounded; profile apparently weakly concavo-convex,
with median ventral fold; periphery of both valves
dorsally geniculated. Length: width ratio 0.53 (ventral
valve), 0.56 (dorsal valve). Ornament unequally
parvicostellate, with 2-3 finer costellae separating
relatively strongly accentuated costellae; very fine
crowded concentric filae are just visible interstitially
between costellae; three faint oblique rugae developed
on posterolateral flanks.
Ventral valve with low apsacline interarea, and
narrow pseudodeltidium extending entire height of
interarea. Interior details of ventral valve unknown.
Dorsal valve interarea very low, orthocline to
weakly anacline, with small, apparently complete
chilidtum. Delicate cardinal process lobes are
continuous laterally with fine, broadly divergent socket
ridges; notothyrial platform beneath cardinal process
is barely thickened above valve floor, extending
anteriorly as a short, low median ridge; transmuscle
septa very poorly developed. Musculature and mantle
canals not deeply impressed; muscle field extends no
more than one-third valve length. Broadly rounded
subperipheral rim slightly raised above dorsal valve
floor, geniculate dorsally in anterior portion; width of
subperipheral rim greatest in posterolateral corner of
valve.
Dimensions
MME 36804 (VV): length 14.8 mm, width 27.7
mm,
MME 36810 (DV): length 17.1 mm, specimen
width 26.8 mm; estimated complete width 30.8 mm;
MME 44968 (DV): length 18.6 mm, width 28.8
mm.
Discussion
Lack of knowledge about interior details of
the ventral valve prevents conclusive identification
of this species as either Platymena or Glyptomena.
In establishing both genera, Cooper (1956, p.882)
commented upon differences between them,
remarking on the flatness of the dorsal valve and
thickened marginal region in Platymena. The delicate
cardinalia and socket ridges, and weakly developed
to barely perceptible septa in the dorsal muscle field
Proc. Linn. Soc. N.S.W., 130, 2009
are more reminiscent of Glyptomena, and although no
dorsal valve exteriors are known for the Gunningbland
species, the sole internal mould seems to suggest a
weakly concave (rather than planar) profile. However,
the presence of a relatively prominent subperipheral
rim is more characteristic of Platymena, to which this
species is tentatively assigned.
Distribution
Gunningbland Formation (upper part) in vicinity
of Gunningbland village, between Parkes and Bogan
Gate, central west NSW; late Eastonian (Ea3-4) i.e.
Katian.
Superfamily Plectambonitoidea Jones, 1928
Family Leptellinidae Ulrich and Cooper, 1936
Subfamily Leptellininae Ulrich and Cooper, 1936
Shlyginia Nikitin and Popov, 1983
Type species: Shlyginia declivis Nikitin and Popov,
1983
Remarks
In addition to describing S. printhiensis from
Molong, NSW, the first species of Shlyginia known
from outside Kazakhstan, Percival (in Percival et al.,
2001) reviewed all six species previously attributed
to this genus. All are similar with respect to general
characteristics of the dorsal valve interior, whereas
there is a wide variation in the size and disposition of
the ventral muscle field. The type species, S. declivis,
has a widely divergent ventral muscle field extending to
about one-third valve length (Nikitin and Popov 1983,
pl. 3, fig. 4; Cocks and Rong 2000, fig. 208, 3b — same
specimen). In Shlyginia fragilis (Rukavishnikova,
1956) the ventral muscle field extends for about one-
third valve length (Rukavishnikova 1956, pl. 2, fig.
18; Popov et al. 2002, pl. 6, figs 22, 25). Shlyginia
extraordinaria (Rukavishnikova, 1956) has a very
large ventral muscle field extending beyond mid valve
length, in which the muscle impressions are conjoined
medially for much of their length (Popov ef al. 2000
pl. 3, fig. 19; Popov and Cocks 2006, pl. 4 figs 22-23).
The ventral muscle field of S. perplexa Nikitin and
Popov, 1996 is much reduced, occupying no more
than one-quarter to one-fifth valve length (Nikitin
and Popov 1996, fig. 4 F-G). The NSW species S.
printhiensis has a ventral muscle scar confined to
the posterior third of the valve, whereas in the new
species described below, the ventral muscle field just
reaches (but never exceeds) half valve length.
Excluded from Shlyginia is S. solida Nikitin and
Popoy, 1984; the sturdy, apparently tubular dorsal
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
median septum of this species indicates that it belongs
in Mabella Klenina, 1984. Also referred to Mabella
on this same criterion is Dulankarella namasensis
Klenina, 1984 (and its synonym D. subquadrata
Klenina, 1984), previously assigned to Shiyginia by
Nikitin and Popov (1996).
Shlyginia rectangularis sp. nov.
Fig. 9 A-X
Diagnosis
Transversely rectangular, dorsoventrally
compressed Shlyginia with distinctive V-shaped
incision at posterolateral extremities of ventral valve;
muscle scar extending to midlength of ventral valve;
2-3 pairs of discrete nodes present on platform of
dorsal valve laterally between muscle field and
peripheral rim.
Etymology
Referring to rectangular outline.
Material
Holotype MMF 44959 (ventral valve); paratypes
include MMF 44947 (conjoined valves), MMF
44948 (ventral valve), MMF 44949 (dorsal valve),
MME 44950 (dorsal valve), MMF 44951 (conjoined
valves), MMF 44952 (ventral valve), MMF 44953
(ventral valve), MMF 44954 (ventral valve), MMF
44955 (dorsal valve), MMF 44956 (dorsal valve),
MME 44957 (ventral valve), MMF 44958 (dorsal
valve), MMF 44960 (dorsal valve), and MMF 44961
(dorsal valve). All specimens are silicified.
Localities
Type locality is L142 (Paling Yards Creek
section at “The Ranch”, Bowan Park), in Quondong
Limestone, Bowan Park Subgroup; also found in same
horizon at L138 (“Quondong”, Bowan Park, east
of Cudal); occurs also at L24 (Licking Hole Creek
area, Walli) in Trilobite Hill Limestone Member of
Vandon Limestone, upper Cliefden Caves Limestone
Subgroup; and at L143 in upper Billabong Creek
Limestone, from outcrop in Billabong Creek at road
crossing, south of Gunningbland [full details of these
localities are given by Percival 1991].
Description
Transversely rectangular shells with long,
straight hingeline, lateral margins nearly straight and
parallel to slightly convergent anteriorly, with broadly
rounded anterior margin. Dorsoventrally compressed,
planoconvex profile; maximum convexity close to
anterior margin; ventral valve flattened medially,
172
becoming broadly sulcate anteromedially in largest
specimens. Valve length between 4.4 and 8.8 mm,
width 5.7 to 13.7 mm; length:width ratios in 13
specimens ranging from 0.55-0.69, with average
of 0.61; maximum width at hingeline with slightly
auriculate, posterolateral extremities in best preserved
specimens, otherwise widest in posterior third of
shell. Ornament finely unequally parvicostellate, very
faintly impressed except for accentuated costellae,
rarely lamellose peripherally in largest specimens.
Ventral interarea low, apsacline, with upper third
of delthyrium covered by small deltidium; dorsal
interarea much lower, anacline, with very fine, paired
chilidial plates flanking trifid cardinal process.
Ventral valve interior: teeth small, unsupported
by dental plates. Muscle field moderately to deeply
impressed, adductor scars confined to a small median
depression deep within delthyrium; diductors much
larger, moderately divergent anteriorly, distinctly
separated medially by fine ridge, and extending to
mid valve length. Mantle canals of lemniscate type,
with moderately strongly impressed vascula media
and weaker vascula genitalia (sometimes not visible).
Narrow, linear median depression extending from
muscle field nearly to anterior margin of valve appears
to exactly coincide with dorsal median septum.
Dorsal valve interior: Anterior edge of hingeline
thickened towards lateral extremities. Cardinalia
typically leptellinine, trifid with prominent central
ridge flanked by finer oblique lateral ridges,
supported on a low thickened notothyrial platform.
Socket ridges short, bladelike and pointed oblique
to hingeline. Muscle field well defined by bounding
ridges extending anteriorly from ends of socket
ridges; muscle field bisected obliquely by low ridges
that may represent proximal traces of vascula media.
Solid ridge-like median septum, not expanding
anteriorly, is separated from front of notothyrial
platform by shallow depression; septum rises sharply
and extends to approximately 0.8-0.85 valve length to
merge with edge of barely undercut platform margin.
Two to three pairs of discrete nodes are present on
platform lateral to muscle field. Mantle canals beyond
muscle field rarely impressed, possibly saccate.
Dimensions
Holotype MMF 44959 (ventral valve) is 7.2
mm long and 11.5 mm wide. Paratype MMF 44947
(conjoined valves) measures 7.4 mm in length, 12.1
mm in width, and 2.0 mm in thickness. Lengths of
12 other paratypes range from 4.4 mm to 8.8 mm,
with most between 5.0-7.5 mm long; widths of 13
paratypes range from 5.7 mm to 13.7 mm, most are
9-12 mm wide. There is no appreciable difference
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Figure 9. Shlyginia rectangularis sp. nov. A— C: exterior of conjoined valves, ventral and dorsal respec-
tively, and posterior profile (dorsal valve uppermost), MMF 44947. D — F: exterior, interior and lateral
profile (posterior to left) of ventral valve, MMF 44952. G — I: exterior, interior and anterior profile of
ventral valve, MMF 44948. J — L: conjoined valves, ventral and dorsal exteriors and posterior profile
(ventral valve uppermost) respectively, MMF 44951. M: interior of ventral valve, MMF 44953. N: inte-
rior of dorsal valve, MMF 44950. O: interior of dorsal valve, MMF 44955. P: interior of ventral valve,
MMF 44954. Q: interior of dorsal valve, MMF 44958. R: interior of juvenile dorsal valve, MMF 44956. S
—T: interior and exterior of dorsal valve, MMF 44960. U: interior of ventral valve, holotype MMF 44959.
V: interior of ventral valve, MMF 44957. W: interior of dorsal valve, MMF 44961. X: interior of dorsal
valve, MMF 44949. Scale bar represents 1 cm. A— C, G —I, N, X from L24, Trilobite Hill Limestone
Member of Vandon Limestone, upper Cliefden Caves Limestone Subgroup at Licking Hole Creek near
Walli; D — F, J—L, M, O, P, R from L135 (east of Copper Mine Creek, near Cliefden Caves), in Trilobite
Hill Limestone Member of Vandon Limestone, upper Cliefden Caves Limestone Subgroup; Q, U, V from
L142 (Paling Yards Creek section at “The Ranch”, Bowan Park), in Quondong Limestone, Bowan Park
Subgroup; S — T from L138 (“Quondong”, Bowan Park, east of Cudal), Quondong Limestone, Bowan
Park Subgroup; W from L143, upper Billabong Creek Limestone at Billabong Creek road crossing
south of Gunningbland.
between measurements of dorsal and ventral valves. shaped incisions at the posterolateral extremities of
the ventral valve interior, and the presence of nodes on
Discussion the platform lateral to the dorsal muscle field — serve
Two distinctive morphological features — the V- to distinguish S. rectangularis from all other known
Proc. Linn. Soc. N.S.W., 130, 2009
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
species of Shlyginia. The function of the V-shaped
incisions is not clear, although one likely explanation
is that they interlock with corresponding thickened
parts of the hingeline in the dorsal valve to strengthen
articulation of the valves when open. Containment of
the dorsal muscle field by bounding ridges is another
characteristic feature of S. rectangularis. The large
ventral muscle field of the new species is comparable
only with that of S. extraordinaria which also has a
similar trapezoidal outline, being noticeably widest
at the hingeline. However, the dorsal interior of S.
extraordinaria, illustrated by Popov et al. (2000, pl. 3,
figs 18-20) and Popov and Cocks (2006, pl. 4, figs 25-
26), exhibits a much less robust median septum than
does S. rectangularis. Unlike both S. extraordinaria
and the other NSW species S. printhiensis, the new
species lacks a well-defined marginal rim in the ventral
valve; the median septum of S. rectangularis is also
relatively much longer than that of S. printhiensis.
Distribution
Limestones of mid-Eastonian (Ea2) age,
equivalent to basal Katian, throughout the Macquarie
Arc in central NSW.
Order Pentamerida Schuchert and Cooper, 1931
Suborder Syntrophiidina Ulrich and Cooper,
1936
Superfamily Camerelloidea Hall and Clarke,
1895
Family Parastrophinidae Schuchert and LeVene,
1929
Parastrophina Schuchert and LeVene, 1929
Type species: Atrypa hemiplicata Hall, 1847
Parastrophina sp.
Fig. 10 A-G
Material
One fragmentary dorsal valve (MMF 44962)
from L147, three ventral valves (all incomplete) MMF
44963-44965 from L24, and one partial ventral valve
(MMF 44966) from L138 (doubtfully attributed).
Localities
Vandon Limestone (Trilobite Hill Limestone
Member), Cliefden Caves Limestone Subgroup at
locality L24, Licking Hole Creek, Walli; Checkers
Member of Regans Creek Limestone at locality
L147, “Red East”, Regans Creek southeast of Cargo;
ventral valve from Quondong Limestone, Bowan
Park Subgroup at locality L138, “Quondong”, Bowan
174
Park, east of Cudal is doubtfully attributed | full details
of localities given by Percival (1991)].
Description
Ventral valve: convex, smooth externally on
posterior and lateral flanks, with shallow sulcus
developed anteriorly, bearing 2-3 costae to form
a weakly plicate anterior margin; internally with
large subparallel dental plates extending to valve
floor, bounding narrow, deep, parallel-sided sessile
spondylium extending to approximately two-thirds
valve length, supported anteriorly by very short
median septum which barely extends beyond anterior
edge of spondylium.
Dorsal valve: smooth, convex posteriorly with
prominent umbo (anterior part of valve not preserved);
cardinal process lacking; deep narrow septalium
present bounded by thin walls anteriorly convergent
on to low thin median septum that extends anteriorly
for an unknown distance; alate plates present.
Dimensions
Dorsal valve MMF 44962 L=
(incomplete), width estimated at 20 mm.
Ventral valve MMF 44963 L=4.5 mm, full width
unknown.
Ventral valve MMF 44966 W= 12.5 mm
(incomplete), estimated width about 20 mm.
7.5 mm
Discussion
The available material, although incomplete, is
assigned to Parastrophina rather than to the externally
similar Camerella on the basis of the presence of
alate plates in the sole dorsal valve. The ventral valve
from the Quondong Formation at Bowan Park has
the same smooth exterior, at least posteriorly, and
similar dimensions to the other specimens. However,
it is only doubtfully attributed to the same species, as
evidence that the spondylium is supported above the
valve floor at the front is lacking (this part of the shell
being broken away). Alternatively, if the dental plates
rest unsupported on the valve floor then this specimen
may be better placed in Stenocamara Cooper, 1956.
Numerous species of Parastrophina have been
described, from North America (Cooper, 1956),
Kazakhstan (Sapelnikov and Rukavishnikova 1975;
Nikitin et al. 1996; Popov et al. 2002; Nikitin et
al. 2006) and elsewhere, but it is difficult to make
accurate comparisons between those (particularly
when described from serial sections) and the sparse
and incomplete silicified material from NSW.
Distribution
Rare in limestones of early Eastonian (Ea2) age,
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
equivalent to earliest Katian, in the Macquarie Arc of
central NSW.
Eoanastrophia Nikiforova and Sapelnikovy, 1973
Type species: Eoanastrophia antiquata Nikiforova
and Sapelnikov, 1973
Eoanastrophia? sp.
Fig. 10 H-I
Material
One specimen, an incomplete dorsal valve,
MMF 44967.
Locality
Quondong Limestone, Bowan Park Subgroup
at locality L138, “Quondong”, Bowan Park, east
of Cudal [full details of locality given by Percival
Geom:
Description
Dorsal valve entirely costate with angular ribs,
occasionally with intercalated costellae; internally
with short septalium supported on long high median
septum; very small sockets; crura present (preserved
only on left-hand side of specimen); no cardinal
process. Ventral valve not available for description.
Dimensions
Specimen 11.4 mm long and 9.4 mm wide (both
dimensions incomplete).
Discussion
Similarly strongly costate parastrophinid genera
include Eoanastrophia Nikiforova and Sapelnikov,
1973 and Maydenella Laurie, 1991. The latter genus,
from the late Middle Ordovician Upper Cashions
Creek Limestone in Tasmania, has a sessile septalium
resting on the valve floor that is bounded by long
subparallel hinge plates, whereas in Eoanastrophia
the hinge plates converge onto a septum which
Figure 10. A — G: Parastrophina sp. A — B: exterior and interior of partial dorsal valve, MMF 44962,
specimen broken during photography; from Checkers Member of Regans Creek Limestone at locality
L147, “Red East”, Regans Creek southeast of Cargo. C — D: interior and exterior of partial ventral valve,
MME 44963; from Vandon Limestone (Trilobite Hill Limestone Member), Cliefden Caves Limestone
Subgroup at locality L24, Licking Hole Creek, Walli. K — G: two interior views (the first slightly tilted to
show dental plates extending to valve floor) and exterior of ventral valve, MMF 44966, doubtfully attrib-
uted to this species; from Quondong Limestone, Bowan Park Subgroup at locality L138, “Quondong”,
Bowan Park, east of Cudal. Scale bar representing 1 cm applies to all specimens in this figure.
H -—I: Eoanastrophia? sp., exterior and interior of dorsal valve (interior view slightly tilted to better show
septum supporting septalium), MMF 44967, from Quondong Limestone, Bowan Park Subgroup at local-
ity L138, “Quondong”, Bowan Park, east of Cudal.
Proc. Linn. Soc. N.S.W., 130, 2009 175
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
supports the septalium (Laurie 1991, p. 85; Carlson
2002, p. 955-958). On this basis, the NSW specimen
is most like Eoanastrophia, although as only one
valve is known, the generic assignment is necessarily
tentative.
Distribution
Presently known only from the one locality in
the Quondong Limestone.
ACKNOWLEDGMENTS
David Barnes (NSW Department of Primary Industries)
expertly prepared the photographic illustrations, and Cheryl
Hormann drafted Figures 1 and 2. Reviews by Robin Cocks
(Natural History Museum, London) and an anonymous
referee greatly facilitated polishing of the manuscript for
publication. Published with the permission of the Director,
Geological Survey of New South Wales, NSW Department
of Primary Industries. This paper is a contribution to
IGCP Project No. 503: Ordovician Palaeogeography and
Palaeoclimate.
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Popov, L.E. and Cocks, L.R.M. (2006). Late Ordovician
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BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
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178
Proc. Linn. Soc. N.S.W., 130, 2009
Rare Fossils (Conulata; Rostroconchia; Nautiloidea) from the
Late Ordovician of Central New South Wales
IAN G. PERCIVAL
Geological Survey of New South Wales, Department of Primary Industries, 947-953 Londonderry Road,
Londonderry, NSW 2753, Australia (ian.percival@dpi.nsw.gov.au).
Percival, I.G. 2009. Rare fossils (Conulata; Rostroconchia; Nautiloidea) from the Late Ordovician of
central New South Wales. Proceedings of the Linnean Society of New South Wales 130, 179-191.
Four decades of detailed palaeontological investigations into highly fossiliferous Upper Ordovician strata
of the Macquarie Arc in central New South Wales has revealed several unique specimens which in some
instances represent the only known examples of phyla or subphyla in this region. Conulariids have not
previously been reported from Ordovician rocks in NSW; here is documented Conularia sp., known
from one specimen found in the Fossil Hill Limestone, and several microscopic specimens of different
genera (including Metaconularia? sp., and the new genus and species Microconularia fragilis) from
deep water allochthonous limestones (Malongulli Formation, and Downderry Limestone Member of the
Ballingoole Limestone). The first Ordovician rostroconch mollusc from NSW is described from a solitary
individual of Eopteria, from the top of the Malongulli Formation. A coiled nautiloid tentatively identified
as Plectoceras from the Gunningbland Formation, again represented by a single specimen, is also described
and illustrated.
Manuscript received 1 December 2008, accepted for publication 16 February 2009.
KEY WORDS: Conulariid, Late Ordovician, Macquarie Arc, Nautiloid, Rostroconch
INTRODUCTION
Rare fossils, often represented by unique
specimens, can sometimes be overlooked in systematic
documentation of a fauna, particularly if they are
not spectacular in appearance or preservation. Yet
such fossils, even when fragmentary or incomplete,
by their very presence can be quite significant
biogeographically. Despite intensive collecting over
more than thirty years (and in some cases around four
decades), the examples described in this paper are the
only known specimens of conulariids, a rostroconch
mollusc, and a genus of tarphyceratid nautiloid that
have been found in Upper Ordovician rocks of the
Macquarie Arc in central New South Wales. Their
uniqueness well qualifies them to be described and
illustrated for the first time.
Stratigraphic setting
The Cliefden Caves Limestone Subgroup and
the overlying Malongulli Formation occur in the
Walli area, between Mandurama and Canowindra in
central NSW (Figure 1). Outcrop of these units has
been mapped in detail south of the Belubula River
by Webby and Packham (1982) in the vicinity of
Cliefden Caves, and by Percival (1976) in the Licking
Hole Creek area, adjoining to the west. Webby
and Packham (1982) established the stratigraphic
nomenclature of the Cliefden Caves Limestone
Subgroup, comprising three formations (in ascending
order: Fossil Hill Limestone, Belubula Limestone,
Vandon Limestone), with the first and last of these
subdivided into a number of members.
Conularia sp. 1s represented by a single specimen
(described herein) collected by G.H. Packham in
the late 1960s from the Taplow Limestone Member
of the Fossil Hill Limestone in the section west
of the “Boonderoo” shearing shed (Webby and
Packham 1982, fig. 3, p.302). No other material of
this species has been found in this or any other level
in the Cliefden Caves Limestone Subgroup, despite
intensive palaeontological investigation of the area
over the past four decades. The age of the Fossil Hill
Limestone is early Eastonian (Eal), equivalent to
latest Sandbian in the middle Late Ordovician. The
Taplow Limestone Member was deposited in shallow
RARE FOSSILS FROM THE LATE ORDOVICIAN
Bogan Gate
L5ty,
a
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SEEDY CREEK
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— sowan PARK
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Figure 1. Locality map showing sites in central New South Wales yielding the Late Ordovician
fossils described in this paper. Outcrop of main Upper Ordovician limestone units shown in black;
localities (L37, L51) in overlying Upper Ordovician clastic-dominated units are shown by spots.
turbulent water interpreted as Benthic Assemblage
(BA) 2 in depth (Percival and Webby 1996); the
conulariid was obtained from skeletal grainstones in
the middle to upper part of the member, overlying
Tetradium cribriforme coral banks.
A solitary outcrop of allochthonous limestone
at the top of the Malongulli Formation on the north-
east flank of Malongulli Trig (Percival 1976) directly
overlies graptolitic shale of early Bolindian age (Bol,
Zone of Climacograptus uncinatus), equivalent
to the latest Katian stage of the Late Ordovician.
A very diverse fauna — including stromatoporoids
(Webby and Morris 1976), radiolaria (Webby and
Blom 1986), sponge spicules (Webby and Trotter
1993), and brachiopods including lingulates (Percival
et al. 1999), strophomenoids and orthoids (Percival
2005), accompanied by numerous fragments of the
nautiloid Bactroceras latisiphonatum Glenister, 1952
(Stait et al. 1985) — is known from acid-processed
180
residues of the limestone. Also present in the residues
are extremely rare conulariid remains, including a
single microscopic conulariid specimen designated
as Microconularia fragilis gen. et sp. nov. and
fragments of a separate conulariid with distinctive
pustulose ornamentation, and a unique specimen of
an articulated rostroconch identified as Eopteria sp.
which, although fragmentary, is recognizable as the
first (and only) known example of this Class in the
Upper Ordovician of NSW. The conulariid material
and the rostroconch are described herein. The
allochthonous limestone is interpreted as having been
initially deposited as periplatformal ooze on the upper
slope (Webby 1992) in BA 4 water depths, prior to
being displaced (after lithification) downslope to its
present BA 5 setting. Thus the conulariids occurring
at this level lived at considerably greater depths than
the larger Conularia sp. from the Taplow Limestone
Member.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
The Bowan Park Limestone Subgroup in the
area east of Cudal (Fig. 1) spans a similar age range
to the Cliefden Caves Limestone Subgroup and
the lower part of the Malongulli Formation, i.e.
early (Eal) to late (Ea 3-4) Eastonian, as indicated
by conodonts studied by Zhen et al. (1999). The
Bowan Park area was mapped in detail by Semeniuk
(1973) who established the internal stratigraphy of
formations and members in use today. One specimen
of Microconularia fragilis gen. et sp. nov. is known
from residues of the Downderry Limestone Member
(of the Ballingoole Limestone at the top of the Bowan
Park Subgroup), which is interpreted as a submarine
channel-fill deposit emplaced at water depths
approximating BA 4 environments. The Downderry
conulariid therefore occupied a comparable habitat to
that of the conspecific example from allochthonous
limestone at the top of the Malongulli Formation.
Upper Ordovician rocks in the Gunningbland
area, west of Parkes on the western side of the
Macquarie Arc (Pickett and Percival 2001), include
the Billabong Creek Limestone (the upper part
of which is correlative with the Cliefden Caves
Limestone Subgroup), and the overlying clastic-
dominated Gunningbland Formation which was
deposited contemporaneously with the Malongulli
Formation, though in slightly lesser water depths.
Faunas of the Gunningbland Formation are dominated
by trilobites (Edgecombe and Webby 2006, 2007)
and brachiopods (Percival 1978, 1979a, 1979b,
2009). Stait et al. (1985) previously described two
coiled nautiloids from this formation, including a
single specimen each of Paradiscoceras dissitum
and an indeterminate tarphyceratid. The specimen of
Plectoceras? sp. described herein is the best preserved
tarphyceratid nautiloid known from this level (a
further fragmentary coiled nautiloid is documented by
illustration only). These specimens, of late Eastonian
(Ea3) age, equivalent to early Katian, are externally
similar to slightly younger tarphyceratids documented
by Percival et al. (2006).
Systematic palaeontology
Type material, comprising specimens described
and illustrated or listed herein, is curated in the
palaeontological collections of the Geological
Survey of New South Wales (designated MMMC for
microfossil specimens, and MMF for macrofossils).
For brevity, authorship of taxonomic hierarchy
above genus level is not cited in the References;
these bibliographic sources are listed in Leme et al.
(2008) for conulariids, Pojeta and Runnegar (1976)
for rostroconchs, and Furnish and Glenister (1964)
for tarphyceratid nautiloids.
Proc. Linn. Soc. N.S.W., 130, 2009
Phylum Cnidaria
Class Scyphozoa Goette, 1887
Subclass Conulata Moore and Harrington, 1956
Order Conulariida Miller and Gurley, 1896
Suborder Conulariina Miller and Gurley, 1896
Family Conulariidae Walcott, 1886
Conularia Miller, in Sowerby 1821
Type species: Conularia quadrisulcata Miller, 1821
Conularia sp.
Fige2
Material
A single incomplete specimen, MMF 44969a-b,
represented by a natural cast and an associated partial
external mould.
Description
The sole specimen includes the upper two-
thirds (approximately) of one individual, extending
25.1 mm in length from the top of the apertural
lobes; maximum width immediately below aperture
is 9.1 mm. Cross-section quadrate, profile steeply
pyramidal with planar to very slightly convex faces
(slightly distorted in preservation) that gently taper
apically, with apical angle estimated to be 8°; apex
not preserved. Ornament consists of narrow, gently
arched transverse ribs (23 per cm) that are defined
by pair of closely-spaced parallel ridges, separated
by interspaces up to three times as broad as the ribs;
interspace ridges barely visible on one face (Fig. 21).
Midline variably expressed, either as a very narrow
ridge (suggestive of an internal carina) across which
the transverse ribs meet in opposition (Fig. 2H), or
a vertical discontinuity across which ribs alternate
(Fig. 21). Transverse ribs are almost everywhere non-
tuberculate except for isolated section of one face (Fig.
2H). Corner sulcus flat-bottomed, with transverse ribs
continuous between adjacent faces. Apertural lobes
triangular in outline and broadly convex in profile,
with continuation of midline; individual lobes are
near vertical in orientation, surrounding a large open
aperture. No internal features preserved.
Discussion
Conulariids are very rare in the Ordovician
of Australia; only a single species has previously
been described from Tasmania by Parfrey (1982),
who established a new genus and _ species,
Tasmanoconularia tuberosa, based on a solitary
partially fragmented specimen (nevertheless with
excellent surface detail) preserved in the Westfield
Sandstone of the Florentine Valley. The brachiopod
181
RARE FOSSILS FROM THE LATE ORDOVICIAN
Proc. Linn. Soc. N.S.W., 130, 2009
182
I.G. PERCIVAL
fauna of this unit (Laurie 1991; age revised by Rong
et al. 1994) contains species of Hirnantia, Kinnella,
Eospirifer, Cryptospira, Onniella? and Isorthis
(Ovalella), which are representative of the Hirnantia
fauna of Hirnantian (latest Ordovician) age.
Parfrey (1982) distinguished Tasmanoconularia
from Conularia and other genera included in the
subfamily Conulariinae Walcott, 1886, by virtue of
the Tasmanian conulariid having a distinct corner
furrow which interrupted the continuity of the
majority of transverse furrows between adjacent
faces. This characteristic suggested affinities with
the Paraconulariinae Sinclair, 1952. Subsequent
opinion (Van Iten and Vyhlasova 2004) and cladistic
analysis (Leme et al. 2008, page 652) has concluded
that Tasmanoconularia is most likely identical with
Conularia, with Leme et al. (2008) advocating that
all previously-proposed families and subfamilies of
conulariids (with the exception of the Conulariidae
Walcott, 1886) be regarded as invalid.
Comparison of C. tuberosa (Parfrey, 1982) with
C. sp. reveals significant differences in size and
ornamentation, sufficient to easily distinguish the two
forms. The Tasmanian species is considerably wider,
attaining a width estimated at 15 mm in an incomplete
specimen, and is much more sharply tapering
towards the apex than is the NSW species. The ribs
of C. tuberosa are crowded together (35-38 per cm)
whereas those of C. sp. are considerably less crowded
(23 per cm). Both species are finely tuberculate, C.
tuberosa conspicuously so; although C. sp. appears
to be almost exclusively devoid of tubercles, this is
most likely an artifact of preservation, as they are
present in one small area (Fig. 2H) of a face that is
less weathered.
Conularia is a long-ranging cosmopolitan genus
with numerous species; furthermore, the cladistic
analysis of Leme et al. (2008) suggests that several
other genera should probably be regarded as synonyms
of Conularia. Comparison of the NSW species with
others assigned to Conularia or its synonyms seems
to be of doubtful value until the genus as a whole is
revised. Coarsely crystalline calcite infilling the sole
specimen of C. sp. has destroyed definitive evidence of
carinae and ridges internal to the corners and midline
(although it is possible the midline is strengthened
internally by a carina — see Fig. 2H). Such features are
significant criteria distinguishing genera and species
of conulariids (Van Iten 1992, Jerre 1994, Leme et
al. 2008), and their absence hinders comparisons with
established taxa.
It is appropriate here to compare C. sp. with
Late Silurian conulariids revised or newly described
from central NSW by Sherwin (1970), as these forms
are closest in age and geography. Mesoconularia
webbyi Sherwin, 1970 has an identical apical angle
of 8° and generally comparable dimensions; however,
transverse ribs on this species are more than twice as
crowded as are those on the Late Ordovician C. sp., and
are always offset across the midline. Paraconularia
packhami Sherwin, 1970, has a very similar apical
angle and spacing of transverse ribs compared to the
older Conularia sp., but in P. packhami the arched
transverse ribs are disjunct and apically depressed at
the midline, whereas in Conularia sp. the transverse
ridges are evenly convex toward the aperture and
may be both continuous and alternating across the
midline.
Distribution
Only known from the Taplow Limestone
Member of the Fossil Hill Limestone, Cliefden Caves
Limestone Subgroup; early Eastonian (Eal) age,
equivalent to latest Sandbian.
Microconularia gen. nov.
Type species (by monotypy): Microconularia
fragilis gen. et sp. nov.
Diagnosis
A microscopic conulariid with non-tuberculate
widely-spaced transverse ribs, lacking a midline;
corners rounded, without furrows.
Discussion
Most Ordovician conulartids are more than 25
mm in length, with only two previously-described
species being less than one-tenth this (Leme et al.
2003, fig. 5). Size would not normally be considered
Figure 2 (LEFT). Conularia sp. A— D: Four faces of internal cast, MMF 44969; E, G, views of corners of
this specimen; F, detail of area of corner outlined on E, showing continuation of transverse ridges across
corner sulcus; H, detail of area of face outlined on B, note minute nodes present on four transverse ridges
adjacent to midline in lower part of enlargement, and continuation of majority of transverse ridges
across midline; I, detail of area of face outlined on D, showing disjunct transverse ridges at midline, and
suggestion of interspace ridges in upper part of enlargement. Scale bar in centre of upper row applies to
A-E and G; scale bar beneath F applies only to the three enlargements. In both instances the scale bar
represents five mm. From Taplow Limestone Member of Fossil Hill Limestone, near Cliefden Caves.
Proc. Linn. Soc. N.S.W., 130, 2009
183
RARE FOSSILS FROM THE LATE ORDOVICIAN
as a distinguishing generic character, but in the case
of conulariids there seems to be a clear dichotomy
between those forms commonly found in inner and
outer shelf environments in a variety of lithologies
(where the overwhelming majority are macrofossils),
and other taxa that are generally known only from
fragmentary remains or microfossils recovered in
acid-insoluble residues of limestones. The latter may
range from relatively shallow to moderate water
depths (BA 2-3), such as those described from Silurian
limestones of central NSW (Bischoff 1973) and the
island of Gotland, Sweden (Jerre 1993), to deep water
(BA 4) settings as interpreted for the forms described
here. One genus is less than 2 mm long, and appears
to be quite distinct from many described macrofossil
conulariids in lacking a definite midline, and in not
developing furrows along the corners. Certainly these
characteristics seem to qualify for differentiation at
genus level, and hence the new genus Microconularia
is proposed.
Teresconularia Leme et al., 2003, from the
Lower Ordovician Santa Victoria Group of the
Cordillera Oriental, northwestern Argentina, shares
with Microconularia the attributes of minute size
(length 1.4 mm) and rounded corners lacking a
sulcus. However, the Argentine genus is considerably
more widely expanding than is Microconularia,
and the latter genus bears much coarser transverse
ribs with strongly angular profiles. The ornament
of Yeresconularia is very fine and crowded by
comparison. There is no evidence of a midline on the
faces of Microconularia, whereas in Teresconularia a
midline is present, albeit very faintly, being marked
by a slight deflection of the otherwise confluent
transverse ribs.
Climacoconus pumilus (Ladd, 1929), most
recently described and illustrated from the Upper
Ordovician Maquoketa Formation of northeastern
Iowa by Van Iten et al. (1996), is another unusually
tiny conulariid up to 2.5 mm in length. Although it
resembles Microconularia in its low apical angle
and coarse transverse ribs, the two genera are readily
distinguished by the pronounced midline and corner
sulcus of C. pumilus.
The maximum length of Eoconularia loculata
(Wiman, 1895), from the Silurian Hemse Beds of
Gotland, is estimated by Jerre (1994) at 10 mm,
approximately 6-7 times as large as Microconularia.
It resembles the new genus in lacking a midline,
and has a similar gradually tapering shape and
coarse transverse ribs. However, the presence of
a corner sulcus in E. /oculata distinguishes it from
Microconularia. The distinctive internal septa of E.
loculata have not been observed in the two known
specimens of the new genus.
184
Microconularia fragilis gen. et sp. nov.
Fig. 3
Diagnosis
As for genus.
Etymology
Genus name in reference to the microscopic size
of the test; species name in reference to the thin and
fragile nature of the specimens.
Material
Holotype MMMC 4388 from L37, allochthonous
limestone at top of Malongulli Formation, head of
Sugarloaf Creek on northeast flank of Malongulli
Trig, near Cliefden Caves; paratype MMMC 4389, a
fragmentary specimen from the same locality as the
holotype; paratype MMMC 4390 from Downderry
Limestone Member of the Ballingoole Limestone,
Bowan Park Subgroup, near Malachis Hill at Bowan
Park.
Description
Test minute, less than 2 mm in length, very
gradually tapering with apical angle of the order
of 1-3°; cross-section quadrate, with flat to slightly
concave faces ornamented with relatively coarse
Figure 3. A—C. Microconularia fragilis gen. et sp.
nov. A — B: Holotype, MMMC 4388, view show-
ing corner, and lateral view of face. C: Paratype
MMMC 4390. Scale bar represents one mm. Holo-
type from locality L37, allochthonous limestone at
top of Malongulli Formation on flank of Malongul-
li Trig; earliest Bolindian (Bol) age. Specimen C
from Downderry Limestone Member of the Ball-
ingoole Limestone, Bowan Park Subgroup, near
Malachis Hill at Bowan Park, late Eastonian (Ea3)
age.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
widely and evenly spaced transverse ribs separated
by interspaces of similar length; 14-19 ribs per mm;
ribs have angular profile (where not eroded) and are
gently and evenly arched; midline lacking. Corners of
test rounded without any furrow; transverse ribs from
adjacent faces are not continuous around corners, but
appear to be offset and alternate so that a rib passes
abruptly into an adjacent interspace. Apertural lobes
and apex not preserved in available specimens.
Internal features unknown.
Dimensions
Holotype MMMC 4388:
maximum width 0.3 mm.
Paratype MMMC 4390: length 1.7 mm, maximum
width 0.4 mm
length 1.5 mm,
Discussion
This exceptionally rare conulariid is represented
in two localities, both in allochthonous limestones
of BA 4 original depositional depth (inferred on
the basis of associated faunas) that have been
redeposited downslope. Age of these horizons is
reasonably contemporaneous (late Eastonian to early
Bolindian).
Distribution
Deepwater strata of late Eastonian (Ea3-4) to
earliest Bolindian (Bol) age, equivalent to Katian
Stage, in central NSW.
Metaconularia Foerste, 1928
Type species: Conularia aspersa Lindstrém, 1884
Metaconularia? sp.
Fig. 4
Figure 4. A— D Metaconularia? sp. A: exterior fragment, MMMC 4391. B: exterior fragment, MMMC
4392. C — D: interior and exterior of fragment, MMMC 4393. Scale bar represents one mm. All speci-
mens from locality L37, allochthonous limestone at top of Malongulli Formation on flank of Malongulli
Trig; earliest Bolindian (Bol) age.
Proc. Linn. Soc. N.S.W., 130, 2009
185
RARE FOSSILS FROM THE LATE ORDOVICIAN
Material
Fragments with tuberculate ornamentation are
uncommon in residues of acid-etched limestones at
locality L37, allochthonous limestone at top of the
Malongulli Formation, head of Sugarloaf Creek on
northeast flank of Malongulli Trig, near Cliefden
Caves. Three representative specimens, MMMC
4391 — 4393, are illustrated.
Description
All material consists of incomplete fragments
of the test, displaying distinctive coarse and fine
tuberculate ornamentation on the exterior surface.
The fragments are flat to gently convex, sometimes
bearing shallow sulci or furrows, and presumably
represent portions of the faces of the theca. Largest
fragment observed is 6.3 mm in length. Tubercles
are irregularly distributed, generally crowded along
shallow furrows in the shell surface (occasionally,
a furrow is underlain by a septum on the interior
surface of the test) and more scattered on the flanks
adjacent to the furrows. A subtle to moderately
strongly expressed longitudinal and _ transverse
arrangement of the tubercles into columns and rows
is often discernable; the rows may be oblique (Fig.
4B) or perpendicular (Fig. 4D) to the main axis of
the specimen. Many (if not all) of the tubercles are
hollow, observable where the tips have been eroded.
Transverse ridges and interrods are lacking, and
sharply defined midlines are not present. Corners
between faces are unknown in the available material.
Internal septa are low narrow linear features, not
twinned; remainder of interior surface is smooth.
Discussion
The tuberculate ornament and absence of
transverse ridges on the faces readily distinguishes
these fragments from specimens of Microconularia
with which they are associated in the acid-etched
residues. Where septa are present internally, their
surficial expression is a crowding of tubercles along
a shallow linear depression or furrow; there is no
development of a deep narrow midline such as is seen
in conulariid fragments from the Silurian age Boree
Creek Formation of central NSW (Bischoff 1978, pl.
1, fig. 1la-b).
Jerre (1993) discussed and figured several
conulariid fragments with comparable tuberculate
ornamentation that he referred to Metaconularia
aspersa (Lindstrém, 1884) from the Silurian of
Gotland. Bischoff (1973) also recognized similar
fragments from both Silurian (Bischoff 1973, pl. 2,
figs. 15 and 17) and Ordovician (pl. 3, fig. 11) horizons,
but did not attribute these to genera. Metaconularia
186
ranges from the Middle and Late Ordovician (Van Iten
and Vyhlasova 2004, fig. 14.1) through the Silurian
(Leme et al. 2008). The specimens from NSW are
too incomplete for definitive identification, so they
are tentatively assigned to Metaconularia pending
collection of more entire material.
Distribution
Recovered only in residues of allochthonous
limestone at top of Malongulli Formation; earliest
Bolindian (Bol) age, equivalent to Katian.
Phylum Mollusca Cuvier, 1797
Class Rostroconchia Pojeta, Runnegar, Morris
and Newell, 1972
Order Conocardioidea Neumayr, 1891
Superfamily Eopterioidea Miller, 1889
Family Eopteriidae Miller, 1889
Eopteria Billings, 1865
Type species: Eopteria typica Billings, 1865
Diagnosis
Eopteriid with a prominent anterior snout that
lacks radial ribs (Pojeta et al. 1977, p. 26).
Eopteria sp.
Fig. 5
Material
Figured specimen MMF 44970a, comprising
a fragmentary pair of silicified conjoined valves.
Several shell fragments definitely attributable to this
specimen were also picked from the same limestone
residue, as was the isolated posterior extremity
MMF 44970b here figured (Fig. 5C). Although the
latter cannot with certainty be assigned to the main
specimen due to absence of intervening shell, there
is a very high probability that it was broken from that
specimen.
Description
Valves moderately biconvex, maximum length
at hingeline; inflated medial third of valves extends
from prominent umbo to ventral margin, and bears
about a dozen rounded radial ribs spaced 2-3 per mm;
raised ribs not present on snout, which instead bears
shallow radial grooves becoming more widely spaced
towards anterior extremity. Posterior third of valves
also marked with shallow radial grooves. Faint closely
spaced concentric growth lines are present on anterior
and posterior flanks. Few internal features visible due
to fragmentary preservation; however, a prominent
Proc. Linn. Soc. N.S.W., 130, 2009
LG.
PERCIVAL
Figure 5. A— C Eopteria sp., conjoined specimen MMF 44970. A: left valve; B: right valve; C: fragment
of posterior, viewed slightly obliquely. Scale bar represents one mm. Specimen from locality L37, alloch-
thonous limestone at top of Malongulli Formation on flank of Malongulli Trig.
internal ridge is present on the interior of the right
valve, trending anteroventrally to intersect the valve
margin. Internal shell surface smooth. Presence of
pegma not verifiable.
Dimensions
The specimen is incomplete, with length of 6.7
mm, and height of 6.0 mm. The separate posterior
extremity is 2.1 mm in length. Estimated maximum
dimensions of the complete individual would be 6-7
mm in height, and at least 9-10 mm in length.
Discussion
This was the specimen from NSW referred to
by Popov et al. (2003, p.177, pers. comm. by I.G.
Percival), in discussion of the palaeogeographic
setting of their new species Eopteria aiteneria from the
Late Ordovician (Hirnantian) Angrensor Formation
of north-eastern central Kazakhstan. That species
was also described from a single damaged shell,
though it is more complete than the NSW specimen
which is of almost identical dimensions. However,
the two are not conspecific, the most significant point
of difference between them being the characteristic
lunulate comarginal ornament developed on the
anterior snout of E. aiteneria. The anterior snout of
the NSW species instead bears several shallow radial
grooves that possibly define a series of flattened wide
ribs progressively decreasing in amplitude away
from the inflated umbo. The medial strongly ribbed
part of E. aiteneria is sharply bounded by carinae,
particularly posteriorly, whereas the NSW species is
more evenly rounded with a relatively gradual change
in slope from the median region to the adjacent flanks.
Proc. Linn. Soc. N.S.W., 130, 2009
The presence of an internal ridge in the right valve of
E. aiteneria cannot be verified as the interior of this
species 1s unknown.
The only other known Late Ordovician species of
Eopteria is E. conocardiformis Pojeta and Runnegar,
1976 from the Little Oak Formation of Alabama
and the High Bridge Group of Kentucky, which is
characterized by an elongation of the anterior snout.
It also appears to be considerably more inflated than
the species from NSW. Cope (2004) assigns an early
Late Ordovician (Sandbian equivalent) age to this
species. Thus Eopteria sp. from NSW, of late Katian
age, 1S significant in partly bridging the gap between
the Laurentian and Kazakhstan occurrences, where
previously no species referable to this genus were
known (Cope 2004, fig. 20). ;
Although Eopteriasp.can bereadily distinguished
from these Late Ordovician (and older) species, it
would be unwise to establish a new species based on
such fragmentary material, and so the specimen is left
in open nomenclature.
Distribution
Recovered only in residues of allochthonous
limestone at top of Malongulli Formation at locality
L37; earliest Bolindian (Bol) age, equivalent to late
Katian.
Class Cephalopoda Cuvier, 1797
Subclass Nautiloidea Agassiz, 1847
Order Tarphycerida Flower, in Flower and
Kummel, 1950
Family Plectoceratidae Hyatt, 1894
187
RARE FOSSILS FROM THE LATE ORDOVICIAN
Genus Plectoceras Hyatt, 1894 Description
Type species: Nautilus jason Billings, 1859 Conch exogastric, planispiral, tightly coiled with
three whorls all in contact, gently expanding from 6-7
Plectoceras? sp. mm height in inner whorls to attain 16.5 mm in height
Fig. 6 at body chamber which is approximately 21 mm in
length; whorl expansion rate (WER) 1.78. Exterior
Material with moderately coarse rounded ribs directed apicad
Specimen MMF 44971, represented by a and forming a wide V-shape at midline; 3-4 ribs in
composite cast, mostly decorticated; this specimen 10mm. Chambers are rounded subquadrate in cross
was longitudinally sectioned for study. section, moderately inflated and gently impressed
Figure 6. A— E Plectoceras? sp., MMF 44971. A— C: conch (prior to sectioning) in lateral view, and two
whorl profiles (slightly rotated) showing ornament and camerae; D — E: longitudinal section through
conch, D off-centre, and E sagittal, with internal features inked-in for clarity. Note in E, remains of sip-
huncle (ventral in position) infilled with calcite immediately above body chamber. F. fragment of exterior
of indeterminate tarphyceratid nautiloid, MMF 44972. Scale bar represents one cm. Both specimens
from Gunningbland Formation at locality L51, “Currajong Park”, Gunningbland.
188 Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
dorsally. Camerae behind body chamber gently flexed
towards aperture and spaced up to 4.3 mm apart,
narrowing to about 2 mm apart in inner whorls where
they extend straight across venter. Siphuncle only
partly visible as a calcite-filled tube with diameter
of 2.4 mm, ventral and marginal in position; septal
necks and connecting rings not preserved.
Dimensions
Maximum diameter of conch 60 mm; width of
body chamber 20.0 mm.
Discussion
Due to its poor internal preservation, with
septal necks absent and the siphuncle inadequately
preserved, identification of this specimen to genus
level remains tentative at this time pending the
discovery of additional better-preserved material. The
ventral position of the siphuncle and strongly ribbed
coiled conch invites comparisons with tarphyceratids.
Of Middle to Late Ordovician genera, the most
similar to the Gunningbland specimen appears to be
Plectoceras, whichis represented by numerous species
in North America. Frey (1995) observed that these
species fell into two major groups, one comprising
forms that are generally smaller in diameter in which
the whorls remain in contact, contrasting with the
second group of generally larger conchs (including
the type species) in which the final whorls became
disjunct. Affinities of the Gunningbland species lie
with the first of these species groups.
The relatively low WER is also similar to that of
Tarphyceras, but that predominantly Early Ordovician
genus is typically nearly smooth externally (B.
Kréger, pers. comm.). Two species of ribbed coiled
nautiloids of latest Ordovician (Hirnantian) age from
the Morkoka River region of the Siberian Platform
were identified as Tarphyceras? by Balashov (1962).
Illustrations of one of these, 7? morkokense Balashov,
1955, clearly show a ventral submarginal siphuncle.
Dimensions of the type specimen (Balashov 1955, pl.
XLII fig. 3a-b; refigured by Balashov 1962, pl. XLVI
fig. 3) are very similar to those of Plectoceras? from
Gunningbland.
Stait et al. (1985, fig. 10) documented an
incomplete external cast of a strongly ribbed coiled
nautiloid from immediately overlying beds in the
Gunningbland Formation, which, in the absence of
any internal features, could only be referred to an
indeterminate tarphyceratid. This specimen is very
possibly congeneric with the one described here as
Plectoceras? as it shares comparable dimensions and
external features. An additional fragmentary exterior
of a similar unidentified coiled nautiloid with coarse
Proc. Linn. Soc. N.S.W., 130, 2009
ribbing is illustrated (Fig. 6F). Slightly younger
tarphyceratids, again with prominent ribs, have been
found in the Jingerangle Formation (of early Bolindian
age, i.e. latest Katian) near Quandialla, about 95 km
south of Gunningbland (Percival et al. 2006), but as all
are preserved as moulds the position of the siphuncle
and other internal features is unknown.
Distribution
Gunningbland Formation (upper part), “Curra-
jong Park” property at Gunningbland; late Eastonian
(Ea3-4), equivalent to early Katian.
ACKNOWLEDGMENTS
Without permission from landholders to collect
specimens from their properties, none of these rare fossils
would ever have come to light; for allowing access I
thank the Dunhill family of “Boonderoo”, the McLarens
of “Liscombe Pools”, and John and Julie Rhodes, former
owners of “Sunnyside”. Technical assistance provided by
Gary Dargan (NSW Department of Primary Industries)
enabled preparation of the polished section of the
tarphyceratid nautiloid from Gunningbland. I am grateful to
Sue Lindsay (Australian Museum, Sydney) for facilitating
SEM imaging of the microconulariids. David Barnes (NSW
DPI) expertly prepared the photographic illustrations, and
Cheryl Hormann (NSW DPI) drafted Figure 1. Heyo Van
Iten, Bjorn Kroger and Leonid Popov provided very helpful
advice on the identifications of the conulariids, nautiloid
and rostroconch respectively. Reviews by two anonymous
referees of the entire manuscript were most useful in
fine-tuning it for publication. This paper is a contribution
to IGCP Project No. 503: Ordovician Palaeogeography
and Palaeoclimate. Published with the permission of the
Director, Geological Survey of New South Wales, NSW
Department of Primary Industries.
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Devonian Marine Invertebrate Fossils from the Port Macquarie
Block, New South Wales
JOHN PickeTT!, Davip OcH* AND EvaAN LEITCH?
1. Geological Survey of New South Wales, NSW, Department of Primary Industries, W.B. Clarke Geoscience
Centre, 947-953 Londonderry Road, Londonderry, NSW 2753, Australia (picketj@bigpond.net.au);
2. Parsons Brinckerhoff, Level 27, Ernst & Young Centre, 680 George Street, Sydney, NSW 2000; GPO Box
5394 Sydney, NSW 2001, Australia (doch@pb.com.au);
3. Department of Environmental Sciences, University of Technology, Sydney, P.O. Box 123, Broadway, NSW
2007, Australia (Evan.Leitch@uts.edu.au).
Pickett, J.W., Och, D.J., and Leitch, E.C. (2009). Devonian marine invertebrate fossils from the Port
Macquarie Block, New South Wales. Proceedings of the Linnean Society of New South Wales 130, 193-
217.
Two assemblages of rugose and tabulate corals, with accessory stromatoporoids and chaetetids, are
described from the Touchwood and Mile Road Formations of the Wauchope — Port Macquarie district
of northeastern New South Wales. Both assemblages are derived from allochthonous limestone clasts,
except that the Mile Road fauna is accompanied at the same level by branching tabulate corals occurring
in the matrix, indicating probable contemporaneity. The fauna from the Touchwood Formation indicates
an Early Devonian (Emsian) age. Macrofossils from the Mile Road Formation indicate a broad Middle
Devonian, probably Givetian age; conodonts accompanying the coral assemblage yield a precise age in
the upper part of the early Givetian varcus Zone. Geographic affinities of the assemblages are typically
eastern Australian, so that if terranes are represented in the block, these were not remote. Stratigraphic and
structural relationships of the units are discussed. The name Mile Road Formation is formally defined.
Manuscript received 27 November 2008, accepted for publication 16 February 2009.
Key words: chaetetids, conodonts, Devonian, Emsian, Givetian, Mile Road Formation, Port Macquarie
Block, Rugosa, stromatoporoids, Tabulata, Touchwood Formation.
INTRODUCTION
Immediately west of Port Macquarie, some
350 km north of Sydney, Palaeozoic units of the
New England Fold Belt are exposed in a series of
narrow belts delimited by NNE-striking faults (Fig.1)
(Leitch, 1980; Roberts et al., 1995). Stratigraphic
relationships between the units and their relative ages
are not clear, so indications of age are particularly
important in geological interpretations of the area.
The ages of these rocks have been little constrained
by published biostratigraphic data, with the only
firm determinations those yielded by conodonts
of Middle-Late Ordovician age from chert in the
structurally dismembered Watonga Formation (Och
et al., 2007), earlier attributed a Silurian or Devonian
age on the basis of meagre conodont and radiolarian
faunas (Ishiga et al., 1988). Unpublished reports
by Pickett (1985, 1991) presented evidence for the
Devonian age of limestone from two other units, the
Touchwood Formation (Leitch, 1980) and the Mile
Road Formation (Taylor, 1984, unpublished; Roberts
et al., 1995). The present article is principally based on
the re-examination of material described by Pickett,
augmented by additional collecting. This has led to
some refinement of the initial results, tectonically
valuable biogeographic information, and a new
stratigraphic interpretation. A formal description of
the Mile Road Formation is included as Appendix 1.
STRATIGRAPHIC UNITS
The Mile Road Formation has only been
recognized in the southern part of the wedge of
rocks bounded by the Sancrox, Cowarra and Sapling
Creek faults (Fig. 1) where it comprises interbedded
fossiliferous siltstone and sandstone, containing
blocks of coralline limestone and silicic tuff. The rocks
form a sequence at least 1500 m thick, dipping steeply
DEVONIAN MARINE INVERTEBRATE FOSSILS
s dncd
Lak¢ Innes (377
rp
485000
Legend
Geological Boundaries
Ue Clarence 4 Mapped Sample Localities
;Moreton - Sc
\ leet Inferred
iy Lithostratigraphic Units HRD
cco
Tablelands
! Recent Sediments Display Boulders at Dam
ee’ aoe al
eee Camden Haven Group
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Fald Belt ~—4 ANY 8 ee iene
A Thrumster Slate Roads
— —— =Faits
[| water ee
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Boulders near Spittway
MRF (Taylor's Locality)
Soh Aushralia
ve
fat os Mingaletta Formation
Watonga Formation A
= Kilometers
6 Os 14 1 ee cas,
Lesa. facia cer 7 —
1 _¥ Port Macquarie Serpentinite
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Figure 1. Geological map of Wauchope — Port Macquarie area.
mostly to the west, and younging in this direction, from northeast away from the Cowarra Fault to almost
based on meagre data from near the intersection of north close to the fault. Sandstone is volcaniclastic
Cowarra Access Road and the Mile Road (GR 478900 and of silicic, probably dacitic, provenance, with
6514400, Grants Head 1:25 000 sheet). Strike ranges abundant detrital plagioclase and vitric and felsitic
194 Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
lithic grains, and uncommon monocrystalline quartz
grains. Many grains are angular and little abraded
suggesting the rocks include abundant little modified
ash. Finer grained rocks are of similar composition.
Scattered coral, brachiopod and crinoid fossils are
locally prominent in the clastic rocks some of which
are extensively bioturbated (cf. Fig. 7C). Coralline
limestone occurs as blocks embedded in fossiliferous
sandstone and siltstone and locally (GR 478400
6514100) as weathered-out boulders up to about 1
m across that may originally have been derived from
autochthonous lenses beyond the outcrop area. Silicic
tuff is prominent in the lower part of the formation
where it forms hard grey beds up to at least 0.25 m
thick. It is of stmilar composition to the sandstones
but distinguished by the presence of well-preserved
shard structures in which the original glass has been
replaced by fine-grained quartzofeldspthic aggregate.
Euhedral plagioclase grains are widespread although
broken angular grains are also common.
The Zouchwood Formation is exposed between
the Lake Innes and Innes Estate faults (Fig.1)
from where it was described by Leitch (1980) as
consisting of a sequence of siltstone, sandstone,
paraconglomerate, basalt breccias and andesite at
least 600 m thick. Much of the stratigraphically
lower sedimentary part of the formation here is thin-
bedded and consists predominantly of simply graded
grey sandstone and darker horizontally laminated
siltstone. Rare paraconglomerate beds, up to at least
20 m thick and the upper part of which are simply
graded, contain clasts of basalt and andesite, slabs
of bedded intraformational material and cobbles of
coralline limestone.
Further west, between the converging Sancrox
and Cowarra faults north of the Mile Road Formation,
Taylor (1984) mapped thin bedded siltstone, some
radiolarian-bearing, graded and massive sandstone,
chert and andesitic breccia as Touchwood Formation.
He considered these rocks were faulted against the
Mile Road Formation, an interpretation followed by
Roberts et al. (1995). The contact between the two
units is unexposed and occurs in a region of very little
outcrop. Although it may be a fault, on the basis of
structural and younging indications in both units, we
favour interpretation as a stratigraphic contact, with
Touchwood overlying the Mile Road (but see below
under Discussion).
Like those of the Mile Road Formation,
Touchwood sandstones are volcaniclastic but differ
in being of more mafic provenance. Abundant
detrital components are lathwork and microlitic lithic
grains and plagioclase; felsitic and vitric grains and
quartz are uncommon, and calcic clinopyroxene is
widespread but mostly only in small amounts.
Proc. Linn. Soc. N.S.W., 130, 2009
FOSSIL LOCALITIES
The fossils described in this article come from
three localities. The first (HRD) lies within the
Touchwood Formation in its type section (Leitch,
1980), the material coming from a disused quarry on
the eastern side of Aston Street, north of the Hibbard
— Port Macquarie road (Hastings River Drive) at GR
490000 6522560 (m), Port Macquarie 1:25,000 sheet
(9435-2S). The material was originally collected by
Erwin Scheibner, and is supplemented by samples
taken by the present authors; Leitch’s (1980, p. 278)
first mention of fossils is restricted to reporting rugose
and tabulate corals. The second (MRF) is within the
informally named “Mile Road Formation” of Taylor
(1984) in a creek-bed in wooded country west of
Forest Road at GR 478400 6514100 (m), Grants
Head 1:25,000 sheet. The material was originally
collected by Michael Taylor, and his formation name
is formalised herein.
Locality MRF could not be re-located using
the information supplied by Taylor, but a general
search led to a third locality (SC) in the bed of Sarahs
Creek south of the ford on an unnamed forestry
track at GR 478500 6514100 on the Grants Head
1:250,000 sheet (9434-1N). Here the mudstones of
the Mile Road Formation dip 65° to 145°, and contain
abundant fragments of the branching tabulate coral
Thamnopora over a stratigraphic interval of possibly
30 m; near the middle of this interval there are also
larger blocks of limestone made up of large colonies
of massive favositid and heliolitid corals, the largest
with maximum dimensions of c. 700 x 450 mm. The
broken fragments of Thamnopora which occur in the
matrix indicate that the larger blocks were derived
penecontemporaneously.
During the construction of Cowarra Damanumber
of large blocks of allochthonous limestone were
uncovered, the largest of which are now on display at
the picnic area near the dam wall. The assemblages in
these blocks indicate that their source is the same as
that of the blocks originally collected by Taylor, but the
assemblages they contain are much richer. In addition
to the small assemblage originally reported by Pickett
(1985) and supplemented herein, the blocks include
large colonies of a large species of Spongophyllum,
Syringopora sp., Heliolites sp., a cystiphyllid, a large
solitary rugosan and Sguameofavosites sp., as well
as brachiopods. Because of the display situation,
none of this material could be collected. The display
boulders were obtained from a locality now covered
by the dam wall at GR 477650 6514250 (Grants
Head sheet), and more material near the spillway at
GR 477950 6514250. All these localities within the
Mile Road Formation are roughly aligned ina WNW
195
DEVONIAN MARINE INVERTEBRATE FOSSILS
— ESE direction, suggesting an episode of slumping
of limy material during deposition of what is probably
the older part of the formation.
The environmental setting of all localities is
similar, in that the fossils are allochthonous, being
derived from clasts in slump deposits. At locality HRD
the limestone clasts are small, the largest observed
being about 35 cm in maximum dimension; some
of the soft-sediment clasts in this deposit exceed a
metre in maximum dimension. All the limestone and
fossil clasts from the Mile Road Formation however
are considerably larger, suggesting that their source
lay much closer than in the case of the Touchwood
Formation.
AGES OF THE OCCURRENCES
The occurrences of coralliform taxa are listed
in Table 1. Those forms only identified in the field
(marked with an asterisk) are not used for age
determination. Detailed discussion is supplied in the
Systematics section, under “Remarks” for each of the
relevant taxa.
Touchwood Formation. Significant for the age of
this unit are Xystriphyllum cf. mitchelli minus, known
only from the mid-Emsian perbonus-gronbergi Zone,
Acanthophyllum sp., whose congeners are restricted to
the Emsian in eastern Australia, and Sterictophyllum
sp., whose genus 1s typically Pragian. Phillipsastrea
Table 1. Occurrences of coral taxa in the Mile Road and Touchwood Formations.
HRD
Ashton St Quarry
Touchwood Fm
MRF SC
Cowarra Dam Sarahs Creek
Mile Road Fm
Mile Road Fm
Chaetetes sp. X
Coenostroma sp. *
Endophyllum cf. columna Hill
Acanthophyllum sp Xx
Xystriphyllum cf. mitchelli minus
Parker =
Phillipsastrea cf. maculosa Hill | x
Sterictophyllum sp. ¥
Favosites salebrosa Etheridge fil.
Pachyfavosites sp.
Squameofavosites squamuliferus
Etheridge fil.
Cladopora sp. X
Thamnopora randsi Jell & Hill
Alveolites sp. A
Alveolites sp. B
Heliolites daintreei group IV Jones
& Hill
=
*Spongophyllum sp.
*Syringopora sp.
*? Squameofavosites sp.
*Heliolites sp.
*cystiphyllid
*large solitary rugosan
Xx
x
Xx
Wstonbor Ne sind :
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
maculosa 1s known from Pragian and Emsian strata,
and the Sguameofavosites squamuliferus group is
typically Early Devonian, although it does range
down into the uppermost Silurian. Some further
slight support for an Emsian age is indicated by the
occurrence of the stromatoporoid Coenostroma. In
summary, the assemblage is taken to indicate a later
Early Devonian age, with a high probability of its
being Emsian.
Mile Road Formation. The coral assemblages
from this unit are less reliably indicative of age than
those of the Touchwood Formation. The best indicator
is probably Endophyllum cf. columna, which suggests
a mid-Givetian age. Thamnopora randsi, on the other
hand, is only known reliably from the mid-Emsian,
whereas Favosites salebrosa is apparently more
typical of Eifelian strata. A small amount of material,
offcuts from the original collection of Michael
Taylor, was digested in acetic acid (Geological
Survey of NSW sample C880), and yielded material
of conodont species which indicate a precise age:
Polygnathus linguiformis klapperi Clausen et al.,
1979, Polygnathus linguiformis weddigei Clausen et
al., 1979, Polygnathus hemiansatus Bultynck, 1987
and Icriodus difficilis Ziegler et al., 1976. The area
of overlap of the ranges of these species, as given by
Bultynck (1987, fig. 9) lies in the upper part of the
lower varcus Zone, of early Givetian age. These taxa
are illustrated in Fig. 2.
DISCUSSION
In the Touchwood Formation the dated material
all occurs as clasts, and in the Mile Road Formation
at least some of the dated material occurs as blocks
embedded in a clastic matrix, and none has been
shown unequivocally to be autochthonous. Thus
the dates provide a maximum age for the units. The
presence of fossils in the matrix as well as in blocks
in the Mile Road Formation suggests the blocks are
penecontemporaneous and hence the Givetian age is
taken as that ofat least part ofthe Mile Road Formation.
For the Touchwood Formation the interpretation is
more equivocal. The limestone here occurs only as
clasts which are restricted to a single bed that is a
debris flow or the product of a high density turbidity
current. Fossils are absent from the surrounding rocks.
There is no record of Devonian limestone clasts in
any of the Carboniferous or Permian units in this
region, and the rocks are of a more mafic provenance
than any of the latter units but similar to those of the
Frasnian Birdwood Formation of the Yarras district
some 25 km further west (Roberts et al., 1995).
This suggests the age of the formation lies within
the Emsian - Frasnian range, and on the basis of its
stratigraphically overlying the Mile Road Formation
can be further restricted to Givetian - Frasnian.
However, the rocks north of Cowarra Dam mapped as
Touchwood Formation have so far yielded no fossils,
and the age suggested by the assemblage from the
type area is older (Emsian) than that of the possibly
underlying Mile Road Formation (early Givetian).
Thus either the assemblage from the Touchwood
formation in its type area does not yield a true age for
the formation or, as originally interpreted by Taylor
(1984), the contact between Mile Road Formation
and Touchwood Formation north of Cowarra Dam is
faulted.
It is noteworthy that the aspect of all fossil
Figure 2. Conodonts from the Mile Road Formation, all x10 except A, x7.5, and E, x20. A—K, Pa elements
of Polygnathus hemiansatus Bultynck, 1987, B and C are oral and oblique views of the same specimen,
E is a juvenile. F, Pa element of Polygnathus linguiformis weddigei Clausen et al., 1979. G, Pa element
of Polygnathus linguiformis klapperi Clausen et al., 1979. H, icriodiform element of Icriodus difficilis
Ziegler et al., 1976.
Proc. Linn. Soc. N.S.W., 130, 2009
7
DEVONIAN MARINE INVERTEBRATE FOSSILS
assemblages is typically Australian. Several taxa
are ascribed to Australian species (Endophyllum
cf. columna, Xystriphyllum cf. mitchelli minus,
Phillipsastrea cf. maculosa, Favosites salebrosa,
Thamnopora randsi), and the Squameofavosites
squamuliferus group is very common in Early
Devonian assemblages throughout eastern Australia.
The genus Sterictophyllum is not known outside
Australia. Thus although the rocks have been
displaced along with the rest of the Hastings Block
(e.g. Cawood and Leitch, 1985) their original location
was well within the Australian province, probably
from a southern continuation of the Tamworth Belt
(Roberts and Geeve, 1999). It is also worth noting
that in the latter a change in sediment provenance
from a region in which intermediate and silicic
volcanism was widespread to one dominated by mafic
volcanic occurred in the Middle Devonian (Cawood,
1983), a change similar to that which occurred
between deposition of the Mile Road and Touchwood
Formations.
SYSTEMATIC PALAEONTOLOGY
The material discussed below is held in the
collections of the Geological Survey of NSW,
indicated by the prefix MMF. Specimens prefixed
AM or AMF are held in the Australian Museum,
Sydney. Literature citations for authors of taxa above
the family level are not cited in the references; they
may be found in Hill (1981).
Phylum PORIFERA Grant, 1836
Class 7?DEMOSPONGIAE Sollas, 1885
Order uncertain
Family CHAETETIDAE Milne-Edwards & Haime,
1850
Genus Chaetetes Fischer von Waldheim in
Eichwald, 1829
Type species
Chaetetes cylindraceus Fischer von Waldheim,
1829.
Remarks
The taxonomy of the fossil group informally
known as chaetetids has been in a state of flux since
the recognition that certain Recent sponges have
a chaetetid morphology, although their spicular
morphology indicates that they are demosponges
(e.g. Ceratoporella: Hartman & Goreau, 1972;
Acanthochaetetes: van Soest, 1984). In the last
overview of chaetetids as a taxonomic group (Hill,
1981) they were regarded as tabulate corals; in recent
treatises on Porifera (Hooper & Van Soest, 2002;
Finks et al., 2004) they have been largely ignored, at
least in terms of updated taxonomy: of the twenty-
nine available generic names given by Hill (1981) in
her review of the “Order” Chaetetida, only two are
mentioned in Finks et al. (2004) and three in Hooper
& Van Soest (2002). On the other hand, modern
genera of “sclerosponges” with chaetetid morphology
receive more exhaustive treatment. It is clear from
Hill’s (1981) introductory remarks that she regarded
the group as polyphyletic, but she has also rendered
the service of bringing together those names relating
to a particular group of morphologies.
In the past, the vertical tubes of chaetetids
have usually been referred to as corallites. In view
of their highly probably sponge nature this seems
inappropriate, so they are here referred to as calicles,
the term favoured for similar features in Ceratoporella
(e.g. Hartman & Goreau, 1972).
Chaetetes sp.
Figure 3 D-G
Material
Two specimens, MMF 32039 and 32040, with
six thin sections. Locality HRD, probably Emsian.
Description
The species forms small, compact masses
reaching at least 5 cm in diameter and 3 cm in
height. The shape appears to have been more or less
hemispherical, but some thin sections show a surface
which bears low mamelons about 7 mm in diameter,
Figure 3 (RIGHT). Spongiomorphs. A-C, topotype specimen of Litophyllum konincki (Etheridge &
Foord, 1894), MMF884, Reid River Limestone, Reid Gap, S of Townsville, Queensland. A, B, transverse
and longitudinal sections, x6; C, detail of B showing vertical trabeculae, x20 approx. D-G, Chaetetes
sp., Touchwood Formation, locality HRD. D, transverse section, MMF32039b, x 3; E, longitudinal sec-
tion of specimen with irregular surface, MMF32040a, x3; F, transverse section, MMF32039a, x10; G,
longitudinal section, MMF32039c, x10. H-K, Coenostroma sp., Touchwood Formation, locality HRD.
H, K, tangential and longitudinal sections, MMF44850, x4.5; J, detail of K showing microstructure of
micropillars, x15.
198
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
whereas others have a smooth surface.
The skeleton is for the most part recrystallised,
but some areas reveal it to have been composed of
fine, near-vertical monacanthine trabeculae about
Proc. Linn. Soc. N.S.W., 130, 2009
0.05 mm in diameter (Fig. 3G). The trabeculae are
united to form walls defining subrounded calicles
about 0.2 mm in internal diameter; wall thickness at
the mid-point is 0.1 — 0.15 mm. In some areas the
199
DEVONIAN MARINE INVERTEBRATE FOSSILS
calicles are interconnected uniserially, but in a rather
meandering pattern (Fig. 3F); in longitudinal section
these appear as pores in the walls. The calicles are
traversed by fine, rather sagging tabulae c. 0.02 mm
thick, generally separated by a distance greater than
the width of the calicle, though this is not always the
case; they number about eleven in 5 mm. The tabulae
display a marked tendency to occur at similar levels
in adjacent calicles, and may even be continuous
through the mural pores.
Remarks
Chaetetids have been reported from Australia in
a number of publications. Etheridge & Foord (1884)
described Amplexopora konincki from Reid Gap,
south of Townsville, north Queensland (Reid River
Limestone, Emsian); Etheridge (1899) reported the
species from Tamworth in NSW and erected for
it the genus Litophyllum. Etheridge’s specimen of
L. konincki from Tamworth (Australian Museum
specimens AM3940, 3941, Moore Creek, near
Tamworth, presumably from the Eifelian Moore
Creek Limestone) is too recrystallised to show
details of wall structure; it is impossible to recognise
whether or not there were trabeculae. It does show
rare connections between calicles. A topotype
specimen (MMF884), rather recrystallised, shows a
microstructure of vertical trabeculae similar to those
of the Port Macquarie material and of other species
of chaetetids (Fig. 3C). However, the tabulae are
crowded (23 in 5 mm) and the spaces between them
always less than the calicle diameter. Connections
between the calicles are rare. I can see no reason for
separating Litophyllum from Chaetetes itself.
Chapman (1918) described Ch. stelliformis
from Early Devonian Loomberah Limestone of the
Tamworth area and, in 1920, Ch. spinuliferus from
an Early Carboniferous Limestone in the Parish of
Mooroowarra, i.e. near Somerton, NSW. Most of
the material reported with this locality information
derives from the hill known as Watts, Babbinboon
(Visean; cf. Campbell, 1957; Pickett, 1967; Moore
and Roberts, 1976), but, in spite of intensive
collecting, the species has not been found there
again. The type specimen in the Museum of Victoria
(P73813, with a longitudinal section; the transverse
section is apparently lost) is clearly a favositid of the
squamuliferus group, revised by Philip (1960), though
not included in his revision; the age of the specimen
is therefore most probably Early Devonian, and the
locality data given by Chapman erroneous, since
there are no outcrops of Early Devonian rocks within
the Parish of Mooroowarra. The species stelliformis
200
is now considered a tabulate coral, Squameofavosites
(Hill, 1950; Philip 1960). Pohler (1998) reported
Pachytheca cf. abdita Yanet, 1972 (in Breyvel’ et al.,
1972) from a stromatoporoid bioherm in the Moore
Creek Limestone Member of the Yarrimie Formation
(Eifelian), but the material was not illustrated or
described. It may be that this is the same form as
Etheridge’s (1899) Litophyllum konincki.
Material of Chaetetes (MMF44896-7) from
the Uglovka Formation in Uglovka quarry, Russia
(upper Serpukhovian) is interesting in that one of
the specimens grew with a smooth surface, while the
other bore abundant mamelons, just as in the Port
Macquarie material, suggesting that this apparent
dimorphism was a regular feature of chaetetids.
Hill (1981) also included desmidoporids and
lichenariids in the order Chaetetida, and the genera
Desmidopora and Lichenaria have both been
reported from Australia (Etheridge, 1902; Fitzgerald,
1955; Hill, 1955, 1957). These occurrences are either
Ordovician or Silurian; they differ considerably from
the present material, and their taxonomic status is not
discussed here.
Class STROMATOPOROIDEA Nicholson and
Murie, 1878
Order SYRINGOSTROMATIDA Bogoyavlenskaya,
1969
Family COENOSTROMATIDAE Waagen and
Wentzel, 1887
Genus Coenostroma Winchell, 1867
Type species
Stromatopora monticulifera Winchell, 1866.
Coenostroma sp.
Figure 3 H-K
Material
MMF44860 from locality HRD.
Description
Specimen fragmentary, but in excess of 32 mm
wide and 9 mm high. Surface apparently smooth and
undulose. Coenostromes dominant, varying widely
in thickness from 0.05 to 0.25 mm 8 — 11 in 2 mm,
separating galleries 0.07 — 0.13 mm high, and which
are subrounded to rather wider than high, consistently
on the same level, generally discrete in longitudinal
section, but occasionally joined laterally over six or
more adjacent galleries. The transverse section shows
a single walled tube 0.6 mm in diameter which may
be a syringoporellid corallite. Coenosteles strongly
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
superimposed, up to 20 observed in a vertical
series, appearing rather meandrine in tangential
section. Microstructure reticulate, of clearly defined
micropillars which are normal to the surface,
appearing as dark spots in tangential section.
Remarks
Coenostroma species do not form a conspicuous
element of eastern Australian Early and Middle
Devonian faunas, as far as they are known (e.g.
Webby et al., 1993; Webby & Zhen, 1993, 1997), the
only published report being Coenostroma sp. from
the Early Emsian (dehiscens Zone) Buchan Caves
Limestone and Heath’s Quarry, Buchan, Victoria
(Webby et al., 1993). The present material differs from
this in its more crowded coenostromes, coenosteles
which are more strongly superposed, and apparently
also in the prominent micropillars of the coenosteles.
Phylum COELENTERATA Frey and Leuckart, 1847
Class ANTHOZOA Ehrenberg, 1834
Subclass RUGOSA Milne Edwards and Haime,
1850
Order STAURIIDA Verrill, 1865
Family ENDOPHYLLIDAE Torley, 1933
Genus Endophyllum Milne-Edwards & Haime,
1851
Type species
Endophyllum bowerbanki Milne-Edwards &
Haime 1951.
Endophyllum cf. columna Hill, 1942a
Figures 4 A-B, 5A
Material
MME2921 2a, 29213a, with two thin sections.
Locality MRF.
Description
Corallum cerioid, exceeding 10 cm in diameter.
Epitheca 0.4 — 0.5 mm thick, showing strong median
dark line. Maximum corallite diameters are 7 — 10
mm. Septa 18 — 22 in each order, the major septa
extending well into the tabularium and sometimes
almost reaching the axis. Minor septa also enter the
tabularium but inside the presepiments are only about
halfas long as the major septa. Even in young corallites
both orders are interrupted peripherally by up to four
rows of steep to almost horizontal presepiments, some
of the inner ones bearing septal crests corresponding
to both orders of septa. Tabulartum 4.5 — 6.0 mm
wide, with tabulae which are flat or slightly concave
near the axis, but turned strongly down and then back
up again in the outer tabularium; 9 or 10 tabulae in
5 mm.
Proc. Linn. Soc. N.S.W., 130, 2009
Remarks
The Queensland species Endophyllum columna
Hill most nearly approaches the present material in
corallite dimensions, though it is generally slightly
larger, in both corallite diameter (10 — 22 mm)
and tabularium diameter (6 — 9 mm), and the wall
thickness is rather less (0.05 — 0.15 mm). Of the
other Australian species of Endophyllum still referred
to that genus, EF. je//li Zhen, 1994 has a much wider
tabularium (10 mm), E. giganteum Zhen & Jell, 1996
has much larger corallites (24—40 mm), and E. banksi
Jell & Hill, 1970a has much larger corallites and more
than twice as many septa.
Endophyllum columna occurs in the upper part
of the Burdekin Formation and the lower beds of the
Cultivation Gully Formation, and is ascribed a mid-
Givetian age by Zhen and Jell (1996).
Family PTENOPHYLLIDAE Wedekind, 1923
Genus Acanthophyllum Dybowski, 1873
Type species
Cyathophyllum heterophyllum Milne-Edwards
& Haime, 1851.
Acanthophyllum sp.
Figure 4C
Material
MMEF32041, locality HRD.
Description
The single specimen is a somewhat oblique thin
section of an eroded corallite near 10 mm in diameter.
In spite of the obliquity of the section the tabularium
appears to be oval rather than round. There are an
estimated 28 major septa; both orders of septa are
thickened in the dissepimentarium, being thickest in
its central part. Near the epitheca they are quite thin.
Minor septa only just reach the tabularium. Septa are
smooth and strongly trabeculate and in their thickest
parts they show a clear zone of trabecular divergence.
Major septa extend almost to the axis; they are
straight in the dissepimentartum but become wavy
in the tabularium. There is a degree of bilaterality of
septa coinciding with the long axis of the section, and
at its margin, on this axis, lies a very short septum,
possibly the counter septum, situated between a
major septum on one side and a minor septum on the
other. The dissepimentarium accounts for about half
the radius of the corallite, the estimated diameter of
the tabularium being 6 mm.
201
DEVONIAN MARINE INVERTEBRATE FOSSILS
Remarks as the distinguishing character between the subgenera
Most acanthophyllids described from Australia Acanthophyllum and Neostringophyllum, although
have a calyx which is either bell-shaped or inverted _ this differentiation has not always been supported
conical. Strusz (1966) took these two calical shapes
202 Proc. Linn. Soc. N.S.W., 130, 2009
{PF PICKEMIS DOCH ANDIE. IE EIRCH
(e.g. Hill, 1981). The only material showing
pronounced fusiform dilatation of the septa in the
dissepimentarium has been referred to the related
species Acanthophyllum clermontense (Etheridge,
1911) and A. kennediense Yu & Jell 1990, both of
which are much larger than the present specimen,
which is not necessarily a fully grown individual.
If the smaller size is a reliable indication, it comes
closest to the material from the Garra Formation
referred to A. aff. clermontense by Strusz (1966).
The Queensland reports of A. clermontense are from
the Emsian (perbonus to inversus Zones; Mawson &
Talent, 2003) Douglas Creek Limestone and the late
Emsian Mount Podge Limestone (Zhen, 1995); A.
kennediense is from the older, Lochkovian to Pragian
Shield Creek Formation (Yu & Jell, 1990). Most of
Struzs’s material from the Garra Formation comes
from the upper levels, so the age is probably late
Emsian (Mawson & Talent, 2000).
Genus Xystriphyllum Hill, 1939
Type species
Cyathophyllum dunstani Etheridge, 1911.
Xystriphyllum cf. mitchelli minus Pedder, 1970a
(in Pedder et al., 1970a)
Figure 4D
Material
MMEF32042, MMF44865 from locality HRD.
Description
One specimen is a small piece of a cerioid colony
which is too thin to permit preparation of a thin
section, but the other has yielded a transverse section.
The weathered surface shows about 20 corallites more
or less in cross section. Corallites range in diameter
from 4.2 mm to 5.8 mm and have 16 — 18 septa in
each order. The major septa reach or almost reach the
axis, but do not appear to interdigitate.
Remarks
In size and septal number the specimen
falls within the ranges of the three smallest
species of Xystriphyllum known from Australia.
Xystriphyllum insigne Hill, 1940a, from Limestone
Siding, Silverwood, Queensland, has diameters in the
range 2 — 4 mm with 12 — 13 septa of each order;
the corallites of X. mitchelli minus Pedder, 1970 (in
Pedder et al., 1970a), from the Taemas Limestone,
Wee Jasper, N.S.W. (mid-Emsian) are less than 6 mm
in diameter, with no more than 20 septa of each order;
and X. parvum Yu & Jell, 1990 has 12 — 15 septa and
diameters of 4— 4.5 mm. Yu & Jell (1990) indicate a
Lochkovian to Pragian age for X. parvum; Mawson
& Talent (1989, fig. 2) suggest an age in the pesavis
— sulcatus Zones, which is in direct agreement with
that of Yu & Jell.
If weight is given to the septal number in
determining the species, then the present material
comes closest to X. mitchelli minus. This form is
known only from the Emsian Taemas Limestone,
from a level within the perbonus-gronbergi Zone
(Pedder et al., 1970a; Mawson & Talent, 2000).
Family PHILLIPSASTREIDAE Hill, 1954
Genus Phillipsastrea d’Orbigny, 1849
Type species
Astrea (Siderastrea) hennahi Lonsdale, 1840.
Phillipastrea cf. maculosa Hill, 1942¢
Figures 4E, 5B
Material
MMF44866, a single fragment from locality
HRD, from which only a longitudinal section could
be prepared.
Description
The slide shows longitudinal sections of one
tabularium of an astraeoid or thamnastraeoid coral, 5
mm in diameter and bounded on either side by strongly
thickened, trabecular fans of a septal stereozone and
its associated ring of horseshoe dissepiments. The
fans are 1.5 — 2.0 mm wide. Septa are robust even in
the outer dissepimentarium, and the dissepimentarial
profile indicates that the everted calyces were raised
Figure 4 (LEFT). Rugose corals from the Touchwood and Mile Road Formations. A, B, Endophyllum cf.
columna Hill, 1942, Mile Road Formation, locality MRF, transverse and oblique longitudinal sections,
MME29213a and 29212a respectively, x1.6. C, Acanthophyllum sp., Touchwood Formation, locality
HRD, oblique section, MMF32041, x3.5. D, Xystriphyllum cf. mitchelli minus Pedder, 1970, Touchwood
Formation, locality HRD, transverse section MMF44865, x4.3. E, Phillipsastrea cf. maculosa Hill, 1942,
Touchwood Formation, locality HRD, longitudinal section, MMF44866, x3. F, G, Sterictophyllum sp.,
Touchwood Formation, locality HRD, transverse and longitudinal sections, MMF44861, x4.5.
Proc. Linn. Soc. N.S.W., 130, 2009
203
DEVONIAN MARINE INVERTEBRATE FOSSILS
Figure 5. Detail of material from Figure 4. A, Endophyllum cf. columna Hill, 1942, Mile Road Formation,
locality MRF, showing details of septa and budding corallite (centre) MMF29213a, x 2.8. B, Phillipsast-
rea cf. maculosa Hill, 1942, Touchwood Formation, locality HRD, MMF44866, showing long major septa
and details of traabecular fans and horseshoe dissepiments, x 8.8.
only a millimetre or so above the general level of the
dissepimentarium. Trabeculae stout, 0.1 — 0.45 mm
in diameter. Tabulae incomplete, the tabularial floor
more or less flat or somewhat raised axially, the rather
confused nature of the section suggesting that the
major septa extend close to the axis.
Remarks
Tabularia are rather larger than those of the type
material (“about 3 mm”), but in its general robustness
the specimen is much closer to P maculosa than
any other Australian species currently referred to
the genus. The tabularia of Bensonastraea praetor
Pedder, 1966 are similar in dimensions, but that genus
has strongly vepreculate septa, of which the present
material gives no indication; the septa of B. praetor
are also less robust than those of the Port Macquarie
specimen. The tabularia of P. carinata Hill, 1942a are
only 3 mm wide and, as the name implies, the septa
are strongly carinate. P. oculoides Hill, 1942d, from
the Garra Formation, has tabularia similar in width
to those of the present specimen, but the septa of
that species are so short that major and minor septa
are of nearly the same length, and the tabulae are
concave or nearly horizontal (see also Wright, 2008).
204
Phillipsastrea currani Etheridge, 1892, as redescribed
by Pedder (in Pedder et al., 1970a), has tabularia up
to 4 mm in diameter, short major septa and horseshoe
dissepiments which are not continuously developed.
Finally, the recently described P. scotti Wright, 2008, is
also close to the present form, but the Port Macquarie
material is too scant for confident attribution to either
this species or P. maculosa.
Phillipsastrea maculosa is known from its type
locality in the Sulcor Limestone (Emsian, serotinus
Zone, Mawson & Talent, 2000), from the Liptrap
Formation at Waratah Bay (Hill, 1954; Emsian,
perbonus-gronbergi Zone, Mawson & Talent, 2000),
the Coopers Creek Limestone at Tyers in Victoria
(Philip, 1962; Pragian, sulcatus to pireneae Zones,
Mawson & Talent, 1994b) and the Late Emsian
(serotinus Zone) Mount Podge Limestone Zhen,
1995). Phillipsastrea scotti is also of serotinus Zone
age.
Suborder CYATHOPHYLLINA Nicholson, 1889
Family CYATHOPHYLLIDAE Dana, 1846
Genus Sterictophyllum Pedder, 1965
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
Type species
Cyathophyllum cresswelli Chapman, 1925.
Sterictophyllum sp.
Figure 4 F-G
Material
A single specimen MMF44861 from locality
HRD, with a transverse and a partial longitudinal
section.
Description
Corallum solitary, apparently cylindrical, with a
maximum diameter of 14.7 mm. Septa long, strongly
radial, of two orders, forming a marginal stereozone
about 3 mm wide, in which the trabeculae are clearly
visible. Septa 25 in each order, the major septa
reaching the axis, where they are carinate; minor
septa long, entering the tabularium. Septa of both
orders taper abruptly after leaving the stereozone.
The imperfect longitudinal section shows no
details of the tabulae, but shows the numerous,
steeply inclined dissepiments inside the stereozone,
and the carinate septa near the axis. The tabulartum
is 5.5 mm wide. Within the stereozone sections of
laterally-growing trabeculae appear as dark spots;
in the inner dissepimentarium they are only slightly
inclined towards the axis.
Remarks
The present specimen is smaller than the
maximum diameters quoted for any of the Australian
species referred to Sterictophyllum, although a single
specimen cannot give any impression of the range of
variation. The stereozone is thicker than in the other
species (S. creswelli (Chapman, 1925) — 2 mm; S.
vallatum Pedder, 1965 —2.5 mm; S. pridianum (Philip,
1962) — 1.5 — 2.5 mm); in S. vallatum, however,
the major septa do not reach the axis. On the basis
of its relative dimensions, the present form appears
to be closest to S. pridianum. (A fourth species,
Mictophyllum trochoides Hill, 1940b, type species of
Cavanophyllum Pedder, 1964, has been included in
the genus by Jell & Hill, 1969, but has major septa
which are somewhat contorted at the axis, lacks the
pronounced stereozone, and is much larger than all
the others. It is not further considered here).
All these species are Early Devonian in age. The
type species (sensu stricto) is known only from its
type locality in the Lilydale Limestone at Lilydale,
Victoria (Pragian, kindlei — pireneae Zones; Mawson
& Talent, 2000); both the other species come from
the Limestone phase of the Coopers Creek Formation
(Pragian, sulcatus to possibly dehiscens Zones;
Mawson & Talent, 2000).
Proc. Linn. Soc. N.S.W., 130, 2009
Subclass TABULATA Milne-Edwards & Haime,
1850
Family FAVOSITIDAE Dana, 1846
Genus Favosites Lamarck, 1816
Type species
Favosites gothlandicus Milne-Edwards
Haime, 1850.
and
Favosites salebrosa Etheridge, 1899
Figure 6 A-B
Synonymy
1899 Favosites basaltica var. salebrosa
Etheridge, p. 166, pl. 21 figs 3-5, pl 27, figs
1-2.
1937 Favosites salebrosa Etheridge; Jones, p.
95, pl. 14, figs 2-6.
1940 Favosites salebrosus Etheridge; Hill and
Jones, p. 197.
2002 Favosites sp. aff. F: salebrosus Etheridge;
Pohler, p. 19, figs SA-D
Material
MME 44857 from the Mile Road Formation,
locality SC.
Description
The material is from fragments ofa large, massive
colony, exceeding 70 x 45 cm in original dimensions.
Corallites range in diameter from 0.65 to 0.83 mm,
with a mean at 0.73. Wall thickness ranges from 0.05
to 0.14, mean 0.06. Mural pores are 0.2 — 0.3 mm in
diameter and at least 0.6 mm apart. Septal spines are
neither conspicuous nor frequent, projecting 0.2 mm
from the wall. There are 14 — 18 complete tabulae in
10 mm.
Remarks
The material accords well with the sections of
the lectotype (AMF 4288. sections AM 47A, B) from
the Woolomol Limestone, in portion 38, parish of
Woolomol, northwest of Tamworth, N.S.W., in which
I have measured rare corallites with a diameter of as
much as 0.9 mm. Jones (1937) reports the species
from what is probably the Cavan Bluff Limestone at
Taemas, without illustration or nomination of material;
this equates to the middle part of the Cavan Formation
of Pedder et al. (1970a), of early Emsian (dehiscens
Zone) age. For the type locality, neither Hill (1942c),
Brown (1942) nor any of the publications of the
Macquarie University group (e.g. Mawson and Talent,
1994a; Pohler, 2001) provides information which helps
age determination. However, this limestone outcrop,
205
DEVONIAN MARINE INVERTEBRATE FOSSILS
a hoe
f
x
hat ol
Figure 6. Tabulate corals from the Touchwood and Mile Road Formations. A, B, Favosites salebrosa
Etheridge, 1899, Mile Road Formation, locality SC, transverse and longitudinal sections, MMF44857,
x5. C, ?Pachyfavosites sp., Touchwood Formation, locality HRD, predominantly longitudinal section,
MMF44863, x4. D, E, Squameofavosites squamuliferus (Etheridge, 1899), transverse and longitudinal
sections, MMF32037, x3.7.
in adjacent portions, is the type locality for some of
the sponges described by Pickett (1969), which are
also characteristic of the lowermost beds of the Timor
Limestone to the southeast. For this interval Pedder
et al. (1970a) have determined an earliest Eifelian
age, so it is probable that the so-called Woolomol
Limestone is more or less coeval. On the other hand,
Pohler (2001, p. 96) indicates that all favositids from
the Tamworth district examined by her are Emsian in
age, and later (Pohler, 2002) describes Favosites aff.
FE. salebrosus from the Emsian Sulcor Limestone, but
206
her material forms cylindrical branches of at least 3
cm diameter, in contrast to the type material, which
is massive, and certainly the present material, which
forms large masses.
Genus Pachyfavosites Sokolov, 1952
Type species
Calamopora polymorpha vat. tuberosa
Goldfuss, 1826.
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
?Pachyfavosites sp.
Figure 6C
Material
A single specimen, MMF 44863, from locality
HRD, with two thin sections.
Description
Corallum massive, original form unknown.
Surface possibly with raised areas. Corallites four to
six sided, 1 — 1.3 mm in diameter. Walls immensely
thickened, so that the lumen diameter is 0.2 — 0.7 mm,
and composed of large bundles of calcite fibres. Mural
pores prominent, about 0.2 mm in diameter. Tabulae
complete, more or less horizontal, irregularly spaced,
possibly reach as many as 14 in 5 mm.
Remarks
This species is much more thickened than the
type, or any other species referred to the genus by
the Russian school, such as P markovskyi (fide
Sokolov, 1962), as the lumen may be all but occluded.
In this respect it is similar to the mature stages of
Riphaeolites Yanet in Sokolov, 1955 (tentatively
included in the family Cleistoporidae by Hill, 1981),
but the present material shows no indication of the
early, less thickened favositid stage characteristic of
that genus .
Pachyfavosites has been reported from Australia
by Pohler (2002), who illustrates both P. rariporosus
Dubatolov, 1963 and P tumulosus Yanet, 1965
from the Emsian Sulcor Limestone Member of the
Yarrimie Formation near Tamworth, NSW; neither of
these species has the intense thickening of the Port
Macquarie material. Riphaeolites is restricted to a
single doubtful record from an unspecified Emsian
limestone (Sulcor?) from the Tamworth area (Pohler,
1998), unaccompanied by either description or
illustration.
Genus Squameofavosites Chernyshev, 1941
Type species
Favosites hemisphericus var. bohemica Pocta,
1902.
Squameofavosites squamuliferus (Etheridge 1899)
Figure 6 D-E
Material
MME32027, with three thin sections; locality
HRD.
Description
The single specimen is a fragment of a cerioid
Proc. Linn. Soc. N.S.W., 130, 2009
colony 30 x 15 mm in diameter and c. 30 mm high.
Corallite diameter ranges from 1.0 to 1.25 mm,
diameters in the lower range being more common.
Wall thickness is variable, from 0.1 to as much as 0.27
mm. Squamulae, though present, are not obvious, the
longest one observed being only 0.1 mm in length.
There are 33 — 40 tabulae per cm. Mural pores have
a diameter close to 0.2 mm, but the preservation is
such that measurements are imprecise. The vertical
distance between their centres is 0.7 — 0.9 mm.
Remarks
Forms which may be referred to the squamuliferus
group in Australia make a fairly homogeneous
assortment (cf. Philip, 1960). The range of variation
described by Philip (1960, notably figs 2, 3) suggests
continuous variation between most of these forms.
All of the material described by Philip (1960),
and the type material of the various taxa involved,
derives from strata of Early Devonian age; in central
western New South Wales the group ranges down into
Late Silurian strata (Pickett & Ingpen, 1990; Pickett
& McClatchie, 1991).
Philip (1960) referred the taxa in this group either
to Squameofavosites grandiporus (Etheridge, 1890) or
to “formae” within Squameofavosites squamuliferus
(Etheridge, 1899), these latter comprising some eight
subspecific units designated by the first eight Greek
letters, and for the first five of which names in the
species category are available (bryani Jones, 1937;
nitidus Chapman, 1914; stelliformis Chapman, 1918;
australis Chapman, 1907; ovatiporus Hill & Jones,
1940). The present specimen fits within the range
reported for forma bryani (Jones, 1937), the type
locality of which is in the Taemas Limestone at Good
Hope, NSW, and which Philip reports from the “Tyers
River Limestone”; both these localities are Emsian,
though the lack of precise localities makes it difficult
to assess the age more accurately.
Family PACHYPORIDAE Gerth, 1921
Genus Cladopora Hall, 1851
Type species
Cladopora seriata Hall, 1851.
Remarks
Following a revision of the type species by
Oliver (1963), Hill (1981) restricts the genus to those
species whose calices are lozenge-shaped in their
mature portion. This definition excludes most of the
Australian species previously included in the genus.
Since the restricted material of this study does not
207
DEVONIAN MARINE INVERTEBRATE FOSSILS
allow the observations necessary for more rigorous
treatment, forms with a consistently rounded calyx
are also treated here.
Cladopora sp.
Figure 7 A-B
Material
MMF 23038, a single fragment of a branching
colony, with one oblique thin section; locality HRD.
Description
The colony is cerioid and branching, the branch
4 mm in diameter. Corallites very small, many in
Figure 7. Tabulate corals from the Touchwood and Mile Road Formations. A, B, Cladopora sp., Touch-
wood Formation, locality HRD, oblique sections of branches. A, MMF32028d, x7.5; B, MMF32038b,
x7.8. C, D, Thamnopora randsi Jell & Hill, 1970. Mile Road Formation, locality SC, random sections,
MMF44858, x2.5. Note bioturbation burrows in C. E, F, Alveolites sp. A., Touchwood Formation, locality
HRD. E, transverse section, MMF32038c, x4.8; F, longitudinal section MMF32038a, x5. G, H, Alveolites
sp. B, Mile Road Formation, locality MRF, portions of large specimen MMF29213a with areas of trans-
verse (G) and longitudinal (H) orientation, x5.
208
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
the cross-section of a branch (at least 18 along a
diameter), rounded in transverse section axially, but
lozenge-shaped towards the margin, 0.35 — 0.4 mm
in maximum diameter, vertical at the axis and curved
gradually towards the surface, which they reach at an
acute angle. Walls thick relative to size of corallites,
without any obvious thickening towards the surface.
Tabulae not observed; there is some indication that
the calices were back-filled with lamellar calcite
rather than by tabulae. Septal spines not observed.
Remarks
Among the Australian species referred to
Cladopora, C. foliata (Jones,1941) is encrusting; C.
gippslandica (Chapman, 1907), originally described
as a bryozoan, and redescribed by Philip (1962),
does not appear to have the lozenge-shaped calices
of the present form; three species from Victoria
(Talent, 1963) (lemaitreae, corrigia, surculus) are
all described as having rounded apertures. I have
examined the type specimen of Cladopora mirabilis
(Etheridge, 1917) (AMF899; 4 thin sections) from
the Reid River Limestone at Reid Gap, south of
Townsville, Queensland. This species forms branches
2 —3 mm in diameter in which the axial corallites are
subrounded, becoming lozenge-shaped only towards
the periphery. In longitudinal section they curve
gently towards the periphery, without geniculation.
There are 6 — 7 corallites along a diameter. Mural
pores are common, tabulae rare, and the wall displays
a prominent median dark line. The present material
differs markedly from C. mirabilis in the much
smaller and more crowded corallites.
Genus Thamnopora Steininger, 1831
Type species
Thamnopora madreporacea Steininger, 1831 (=
Alveolites cervicornis de Blainville, 1830).
Thamnopora randsi Jell and Hill, 1970b
Figure 7 C-D
Material
Two large blocks of mudstone containing
abundantly branching coralla, MMF 44858 (2 thin
sections), MMF 44869, Mile Road Formation,
locality SC.
Description
Corallum branching, bifurcating, branches
cylindrical, 6 — 14 mm in diameter, mature branches
being generally in the upper range. In transverse
section there are 8 — 10 corallites along the median
Proc. Linn. Soc. N.S.W., 130, 2009
plane ofa mature branch. Diameter of mature corallites
1.2 — 1.5 mm, their combined walls about 0.2 mm in
thickness near the axis, and 0.6 — 0.8 mm in the outer
stereozone, which is 2.5 — 3 mm wide. In the inner
parts of the branches the walls show a conspicuous
median dark line, but this becomes much more diffuse
in the outer stereozone, where the stereome may show
a lamination parallel to the surface. Near the axis the
corallites are parallel to the branch, but turn outwards
without geniculation to reach the sides of the branch
at about 45°. Mural pores have a diameter of 0.2 — 0.3
mm, and are rather funnel-shaped in the stereozone.
They occur in a single series on the faces of the
corallites. Calices are 3 — 4 mm deep. Septal spines
are rare, < 0.1 mm in length, conical, not trabeculate,
and occur both at the axis and in the stereozone. Latex
replicas from natural moulds show no sign of septal
ridges in the calyces. Tabulae are complete, lie closer
together than the width of the corallite, usually 4 in
2 mm.
Remarks
The present material accords well with that
described by Jell and Hill (1970b), though the branches
are slightly thinner (14 mm as against 15 mm in the
types), and the corallites open slightly more obliquely
to the sides of the branches. A significant similarity is
the way the median dark line becomes less obvious
towards the exterior, and the presence of growth
lamination in these areas. Thamnopora plumosa Jones,
1941 has much stouter branches with nearly twice
as many corallites across the median plane, and the
thickening is less conspicuous. Thamnopora foliata
Jones, 1941 is laminate; 7: meridionalis (Nicholson
and Etheridge, 1879) is more delicately branched;
T. crummeri (Etheridge, 1899) is closer, but has fewer
corallites across the median plane and the difference
in the amount of thickening between the axial and
outer zones is less pronounced (I have examined the
sections of the holotype, AM 3981 and 4687). The
Victorian species, T alterivalis (Chapman, 1914),
T. angulata Hill, 1950 and 7 tumulosa Hill, 1950
all have significantly thinner branches, the largest
reaching only 7 mm.
Thamnopora randsi 1s known so far only from its
type area, the Douglas Creek Limestone, of Clermont,
Queensland (mid-Emsian, perbonus to inversus
Zones; Mawson & Talent, 2003).
Family ALVEOLITIDAE Duncan, 1872
Genus Alveolites Lamarck, 1801
209
DEVONIAN MARINE INVERTEBRATE FOSSILS
Type species
Alveolites suborbicularis Lamarck, 1801.
Alveolites sp. A
Figure 7 E-F
Material
Four indifferently preserved — specimens,
MMEF32038, 44862, 44867 and 44868, with six thin
sections, all from locality HRD.
Description
All the material is of small fragments, the largest
being less than 2 cm in maximum diameter. Corallites
usually crescentic, apparently reclined, up to 0.4 mm
high and 0.8 — 1.0 mm wide. Squamulae, septal spines
and mural pores not observed. Tabulae 0.25 — 0.4 mm
apart.
Alveolites sp. B
Figure 7 G-H
Material
Three specimens, MMF 29213 — 29315, with
three thin sections, all from locality MRF.
Description
Corallum moderately large, exceeding 10 cm in
maximum dimension. Corallites generally crescentic,
but occasionally polygonal, 0.3 — 0.5 mm high and
0.5 — 0.7 mm wide. Occasional short septal spines
occur on the side of the corallite away from the curved
surface. The wall thickness varies considerably
within the corallum, some areas having corallites
whose walls reach only 0.02 mm, but for the most
part the walls are thickened, reaching 0.1 or even 0.2
mm in extreme cases. The non-thickened areas pass
rather abruptly into thickened areas, and individual
corallites may lie partly in each of the two. There are
occasional mural pores. Tabulae are at right angles to
the direction of growth of the corallite, thin even in
the thickened parts of the corallum, complete, and 0.2
— 0.6 mm apart.
Remarks
For all that the Australian literature refers to
some twenty species of A/veolites (Pickett, 1999), the
genus is not well documented in this country, either
morphologically or stratigraphically. Six species
(caudatus Hill, 1954; intermixtus Lecompte, 1939;
multiperforatus Salée, 1916 (in Lecompte, 1933);
saleei Lecompte, 1933; swborbicularis Lamarck,
1801; tmida Hinde, 1890) are known from Late
Devonian strata in Western Australia. A further two
were established by de Koninck (1876; obscurus,
rapa) and, as the type material was destroyed by
fire and details of the type localities are vague, it is
probably better that the names be allowed to languish;
his other three reports are unillustrated and, based as
they are on external features alone, should be regarded
as dubious. Chapman’s (1921) species regu/aris and
victoriae were regarded as species of Favosites by
Philip (1960).
The status of the remaining eight taxa is not
necessarily sound. The holotype of the only Silurian
species, A/veolites piriformalis Etheridge, 1921 from
the Yass district, has never been traced, and its internal
structure is inadequately known. The holotype of 4.
queenslandensis Etheridge & Foord, 1884, from the
Emsian Reid River Limestone at Reid Gap, south of
Townsville, has not been traced and the species has
not been redescribed since the original publication.
Hill et al. (1967) illustrated, without description,
forms referred to A. sp. ex gr. fecundus (Salée, 1916)
and A. sp. nov. aff. /emniscus Smith, 1933, of which
the first has a branching corallum and the second
does not show the areas of thin- and thick-walled
corallites of A. sp. B. from locality MRF. A/veolites
stamineus Hill, 1950 from the Emsian Murrindal
Limestone at Buchan, Victoria, is a distinctive, thinly
encrusting form. Neither of the forms referred to
A, suborbicularis Lamarck, 1801 or A. sp. nov. aff.
A. hemisphericus (Chernyshev, 1937) by Brihl &
Pohler (1999) shows areas of thin- and thick-walled
corallites, apart from the thinner-walled basal layer
of A. suborbicularis. Finally, the material referred to
A. sp. aff. A. taenioformis Schliiter, 1899 by Philip
(1962) forms encrusting layers no more than 4 mm
in thickness.
The material described here as A/veolites sp. A is
too scant for proper identification, and that described
as Alveolites sp. B does not appear to be the same as
any Australian forms so far reported.
Order HELIOLITIDA Frech, 1897
Family HELIOLITIDAE Lindstrém, 1876
Genus Heliolites Dana, 1846
Type species
Astraea porosa Goldfuss, 1826.
Heliolites daintreei Nicholson & Etheridge, 1879
group IV Jones & Hill, 1940
Figure 8
Material
Two specimens, fragments of much_ larger
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
Figure 8. Tabulate corals from the Mile Road Formation, locality SC. A, B, Heliolites daintreei Nicholson
& Etheridge, 1879, group IV Jones & Hill, 1940. A, transverse section MMF44856, x5.3; B, longitudinal
section MMF44859, x5.7.
colonies, MMF 44856, 44859, from locality Sarahs
Creek.
Description
Corallum massive, large. Tabularia consistently
1.2 mm in diameter, but ranging up to 1.5 mm. No
areola is developed, but the tabularia are surrounded
by 19 —20 tubules of varying size; tubules throughout
the coenenchyme range from 0.2 to 0.5 mm in
diameter. Tabularia separated by 3 — 10 tubules. Septa
12, laminar, apparently without axial spines, reaching
about halfway to the axis. There are thin horizontal
zones in which the skeleton is slightly thicker; these
are about 2 mm thick and 7 — 9 mm apart. The tabulae
are 0.6 — 1.1 mm apart, and the diaphragms about 11
in 5 mm.
Proc. Linn. Soc. N.S.W., 130, 2009
Remarks
Since the review of Australian Silurian and
Devonian heliolitids by Jones and Hill (1940) no other
overview of the group has been attempted. There is a
clear need for any proper study of the group to be
based on extensive material, allowing population
studies. Here we simply follow the work of Jones and
Hill.
Heliolites daintreei, as conceived by Jones
and Hill (1940), is an enormously variable species
ranging from the Late Silurian to the Early Devonian,
even the four informal “groups” not demonstrating
reliable stratigraphic range. The present material, as
it appears to lack axial spines on the septa, does not fit
comfortably in any of the taxa recognised by them.
Dil
DEVONIAN MARINE INVERTEBRATE FOSSILS
ACKNOWLEDGMENTS
We thank Mike Neville of the NSW Department of
Commerce for information on the source of the limestone
olistoliths on display at Cowarra Dam. David Barnes and
Yong-yi Zhen helped with photography and plate assembly.
JWP is particularly grateful to Ruth Mawson for her
generous help with the conodont determinations. Michael
Taylor originally discovered the limestone fossils in the
Mile Road Formation and first recognized the Touchwood
Formation west of the Cowarra Fault. The Birpai Aboriginal
Land Council graciously granted permission to collect
material on their land.
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216
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
APPENDIX 1.
Mile Road Formation
Definition: The name Mile Road Formation is
applied to interbedded fossiliferous siltstone and
sandstone, containing blocks of coralline limestone
and possibly autochthonous limestone lenses, and
silicic tuff, mapped as stratigraphically underlying
the Touchwood Formation in the eastern part of the
Hastings Block.
Synonymy: The unit was first recognised by Taylor
(1984) who termed it the Mile Road Formation. The
unit was referred to as the Mile Road beds by Roberts
et al. (1995).
Derivation of name: Named from the Mile Road that
traverses part of the unit in the Cowarra State Forest
(GR 478700 6514700 Grants Head 1:25 000 sheet).
Distribution: The Mile Road Formation is known
only from the eastern Hastings Block where it has
been recognised in the southern part of a slender
wedge bounded by the Cowarra, Sapling Creek and
Sancrox faults. It is mapped over an area of about 7
km’.
Type section: Neither Taylor (1984) nor Roberts
et al. (1995) designated a type section although the
latter authors specified a type locality on the Cowarra
Access Road (GR 479000 6514800 to 478900
6513100, Grants Head 1:25 000 sheet). This locality
lies nearly along strike and encompasses only the
lower part of the formation. We suggest that the type
section be that extending northwest from the Cowarra
Access Road at GR 478800 6513800 (base) along a
tributary of Sarah Creek to GR 478300 6514300 (top)
(Grants Head 1:25 000 sheet).
Stratigraphic relationships: Neither base nor top
of the unit is exposed. It is truncated downwards
by the Cowarra Fault and is here interpreted as
being stratigraphically overlain by the Touchwood
Formation | — 2 km south of the Oxley Highway.
Thickness: A maximum preserved thickness of
between 1500 and 2000 m is estimated based on the
mapped width of the unit and the assumption of an
overall steep northwest dip and consistent northwest
direction of younging.
Content: Medium to thick bedded volcaniclastic
siltstone and sandstone of intermediate-silicic
Proc. Linn. Soc. N.S.W., 130, 2009
provenance, locally bioturbated and/or fossiliferous
with crinoids, brachiopods and corals. Widespread
breccias/conglomerates with coralline limestone
clasts to c. | m set in a coarse sandy matrix. Grey
hard massive silicic tuff interstratified with epiclastic
rocks.
Age and correlation: A small conodont assemblage
from probably penecontemporaneously derived
allochthonous blocks gives a precise age of the upper
partofthelowervarcus Zone, early Givetian. Significant
taxa are Polygnathus linguiformis klapperi Clausen et
al., 1979, Polygnathus linguiformis weddigei Clausen
et al., 1979, Polygnathus hemiansatus Bultynck, 1987
and Icriodus difficilis Ziegler et al., 1976. Additionally
the blocks contain an abundant macrofauna of rugose
and tabulate corals, spongiomorphs and brachiopods;
the branching tabulate coral Thamnopora, occurring
in the bioturbated matrix, suggests strongly that
the blocks are penecontemporaneous, and that the
conodont assemblage indicates a real age, at least for
that part of the Formation.
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Age Determination and Growth in the Male South African Fur
Seal Arctocephalus pusillus pusillus (Pinnipedia: Otariidae)
Using External Body Measurements
C. L. Stewarpson’, T. Prvan*, M. A. MEYER? AND R. J. RitcHie**
'Botany and Zoology, Australian National University, Canberra, ACT, Australia. (Present address, Fisheries
and Marine Sciences Program Bureau of Rural Sciences, The Department of Agriculture, Fisheries and
Forestry, CANBERRA ACT 2601 Australia).
"Department of Statistics, Macquarie University, NSW 2109, Australia.
*Marine and Coastal Management (MCM), Rogge Bay, Cape Town, South Africa.
*School of Biological Sciences, The University of Sydney, NSW 2006, Australia.
*Corresponding Author (rrit3 143 @usyd.edu.au)
Stewardson, C.L., Prvan, T., Meyer, M.A. and Ritchie, R.J. (2009). Age determination and growth in the
male South African fur seal Arctocephalus pusillus pusillus (Pinnipedia: Otariidae) using external body
measurements. Proceedings of the Linnean Society of New SouthWales 130, 219-244.
Morphology, relative size and growth of the South African fur seal or Cape fur seal, Arctocephalus
pusillus pusillus, from the coast of southern Africa are described and comparisons made to data available
on the closely related Australian fur seal (Arctocephalus pusillus doriferus) and the New Zealand fur seal
(Arctocephalus forsteri). Useful information can be gained from body measurements of seal carcasses
provided canine teeth are extracted for aging. External body measurements (12 linear variables) were
examined in relation to standard body length (SBL) and chronological age (y) using linear regression and
non-linear least squares fitting as appropriate. Animals ranged from < 1 month to > 12 y. Of the 149 animals
in the study, 39 were animals of known-age based on tagging; 34 were aged from highly reproducible
counts of incremental lines observed in the dentine of upper canines (i.e., range 1-10 y); 10 were identified
as adults > 12 y (1.e., pulp cavity of the upper canine closed); and 66 were not aged. At birth, male South
African fur seals are 35% (c. 69 cm) of their mean adult size. At puberty, they are 57% (c. 113 cm). The
foreflippers measure 25—26% (c. 18 cm) of standard body length (SBL) in pups, and 24% (c. 48 cm) of SBL
in adults. The hind flippers are considerably shorter, measuring 19% (c. 13 cm) in pups, and 14.5% (ce. 29
cm) in adults. Axillary girth is usually about 57-67% of SBL. Growth of SBL was rapid during the early
postnatal period with a significant growth spurt occurring at the onset of puberty (2-3 y). The rate of growth
slowed significantly between 6 and 7 y. Social maturity was reached at about 9 to 10 y. Growth slowed
thereafter. The mean SBL for aged males >10 y and unaged animals > 200 cm was 199 cm. Relative to
SBL, facial variables and the fore/hind limbs scaled with negative slope relative to SBL or were negatively
allometric; tip of snout to genital opening scaled with positive slope; and tip of snout to anterior insertion of
the foreflipper was positively allometric. Relative to age, body variables scaled were negatively allometric.
SBL was found to be a ‘rough indicator’ of age and age group. The growth kinetics of juvenile and adult the
South African fur seal and the Australian fur seal are best described by the logistic and double exponential
(Gompertz) models rather than the exponential von Bertalanffy model. Australian fur seals grow at a faster
rate but asymptotic maximum sizes are similar in South African and Australian fur seals.
Manuscript received 21 May 2008, accepted for publication 17 December 2008.
Key words: Allometry, Arctocephalus pusillus doriferus, Arctocephalus pusillus pusillus, Australian fur
seal, body, growth, growth curve modelling, pinnipeds, South African fur seal.
evolutionary links within and between populations of
INTRODUCTION the same species and between species. Growth and
body-size estimates can be used for monitoring the
Data on the physical growth of pinnipeds is effects of population pressures and changes in the
important to understanding the biology, ecology and quality of the habitat of marine mammals (Bester
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
and Van Jaarsveld, 1994). Within the Otariidae (fur
seals and sea lions) quantitative descriptions of
growth in body length based on animals aged from
tooth structure, or on animals of known-age (1.e.,
animals tagged or branded as pups), are available for
several species of fur seals and sea lions including the
Australian fur seal (Arctocephalus pusillus doriferus)
(Arnould and Warneke, 2002) which is very closely
related to the South African fur seal (Wynen et al.,
2001); the New Zealand fur seal (Arctocephalus
forsteri) (Dickie and Dawson, 2003; McKenzie et
al., 2007), the subantarctic fur seal (Arctocephalus
tropicalis) (Bester and Van Jaarsveld, 1994), the
Antarctic fur seal (Arctocephalus gazella) (Payne,
1979; Krylov and Popov, 1980; McLaren, 1993), the
Northern fur seal (Callorhinus ursinus) (Scheffer and
Wilke, 1953; Bychkov, 1971; Bigg, 1979; Lander,
1979; McLaren, 1993; Trites and Bigg, 1992, 1996)
and the sea lions, Eumetopias jubatus, the Steller sea
lion (Fiscus, 1961; Thorsteinson and Lensink, 1962;
Calkins and Pitcher, 1983; Loughlin and Nelson,
1986; McLaren, 1993; Winship et al., 2001), and
Otaria byronia, the South American sea lion (Rosas
endle 1993):
Physical growth in the northern fur seal and Steller
sea lion have been studied in the most detail and is
based on the largest number of animals of known age.
The general growth curve for the Northern fur seal
and the Steller sea lion is presumably representative
of all highly polygynous male otartids. Male pups
of Northern sea lions measure c. 66 cm at birth and
grow at a steady rate (Scheffer and Wilke, 1953;
Trites and Bigg, 1992, 1996). Growth is claimed to
increase suddenly at 3-4 y (puberty) and slows soon
after attamment of social maturity (McLaren, 1993).
Estimated asymptotic length is about 189 cm for
males > 4 y, and is reached by c. 12 y in most animals
(McLaren, 1993). Growth curves of the Steller sea
lion are basically similar in shape and also claimed
to best fit a logistic rather than exponential saturation
curve (Winship et al., 2001). Asymptotic maximum
size of the Steller sea lon is much larger than fur
seals: maximum size of males is about 3 m and 700
kg at about 12 y.
The limited information on growth in body
size available for South African fur seals was based
on measurements that were aged physiologically
(cranial suture age) rather than chronologically (y)
(Rand, 1956). Unfortunately, in South African fur
seals cranial sutures are not a very reliable guide to
age (Stewardson, 2001; Stewardson et al., 2008).
Comparisons will be made to data available on the
Australian fur seal (Arnould and Warneke, 2002),
the New Zealand fur seal (Dickie and Dawson, 2003;
220
McKenzie et al., 2007) and the subantarctic fur seal
(Bester and Van Jaarsveld, 1994). Apart from studies
by Scheffer and Wilke (1953) and Payne (1979)
information on the relative growth of external body
measurements of other fur seals is scant, e.g., axillary
girth vs. standard body length, length of limbs vs.
standard body length.
Here we examine the body measurements of 149
male South African fur seals, Arctocephalus pusillus
pusillus, from Southern Africa. Specific objectives
were to: (i) describe the general morphology of the
animal; (11) quantify growth of body measurements
(12 variables) relative to standard body length (n =
134 animals) and chronological age (7 = 83 animals),
(iii) determine if standard body length (SBL) is a
useful indicator of age, (iv) compare three commonly
used models for the growth kinetics of South African
fur seals compared to Australian fur seals (exponential
saturation curve or von Bertalanffy curve, Logistic
curve and the double exponential or Gompertz curve)
(Zullinger et al., 1984; Zeide, 1993).
MATERIALS AND METHODS
Collection of specimens
South African fur seals were collected along
the Eastern Cape coast of South Africa between
Plettenberg Bay (34° 03’S, 23° 24’E) and East
London (33° 03’S, 27° 54’E), from August 1978
to December 1995, and accessioned at the Port
Elizabeth Museum (PEM). From this collection, 110
males were selected for examination. Apart from
specimens collected before May 1992 (n = 38), all
specimens were collected by the first author. PEM
animals were aged based on dentition (n = 32), some
PEM animals were aged using dentition growth rings,
animals designated >12 y (n = 10) were animals with
12 growth rings in their teeth but their pulp cavities
were closed and so no more growth rings could be
deposited and so were at least 12 y old but could have
been older. One animal (PEM2238) was collected NE
of the study area, at Durban.
Measurements from 39 males from Marine
and Coastal Management (MCM), Department
of Environment Affairs and Tourism, Cape Town
were also available. These measurements were
from animals that had been tagged as pups, and
were therefore of known-age (1-13 y). MCM seal
specimens are accessioned as MCM followed by a
number. The accession numbers of all the animals
used in the present study are listed in Appendix 1.
The full data set is accessible in the public domain
(Stewardson, 2001).
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
Body measurements
Standard necropsies were performed and biolog-
ical parameters recorded, based on recommendations
of the Committee on Marine Mammals, American
Society of Mammalogists (1967). Upper canines were
collected for age determination. The skull is probably
the most useful part of a seal carcass to retain for
later study but it is not always possible to arrange
for the skull of a dead seal to be retained. Nuisance
seals are sometimes culled to satisfy the concerns
of aquaculture and fisheries interests. From humane
considerations, permits for such culls usually specify
that the animals are fatally shot in the head, which
ruins the skulls for morphological studies, but teeth
for aging can usually be retrieved (Thorsteinson and
Lensink, 1962; Pemperton et al., 1993; Winship et al.,
2001; Arnould and Warneke, 2002; McKenzie et al.,
2007). Body measurements of seal carcasses are most
useful if canine teeth are extracted for aging.
Measurements (12 variables) were taken to the
nearest 5 mm (0.5 cm) using a flexible tape measure or
vernier callipers as appropriate (Figure 1). Although
body weight and blubber thickness were recorded,
these measurements were not included in the analysis
because they can vary according to physiological
condition, e.g., body condition is influenced by
seasonal fluctuations in food supply, illness or injury,
and breeding condition. The blubber of Australian fur
seals is known to vary seasonally with a maximum
in late austral spring (Arnould and Warneke, 2002).
Apart from specimens collected before May 1992, all
PEM measurements were recorded by the first author.
The majority of MCM measurements were recorded
by the third author.
Age determination
The age of animals was estimated from counts of
Growth Layer Groups (GLGs) observed in the dentine
of thin tooth sections (Payne, 1978; Oosthuizen,
1997; Oosthuizen and Bester, 1997; Stewardson et al.,
2008). Upper canines were sectioned longitudinally
using a circular diamond saw. Sections were ground
down to 280-320 pm, dehydrated, embedded in resin
and viewed under a stereomicroscope in polarised
light (Oosthuizen, 1997; Oosthuizen and Bester,
1997). Each section was read by one individual five
times, without knowledge of which animal was being
examined (repeated blind counts) similar to Payne
(1978). Ages were rounded off to the nearest birth
date. The median date of birth was assumed to be |
December (Shaughnessy and Best, 1975), which is
similar to the mean date of birth for Antarctic fur seals
(Payne, 1978). The median of the five readings was
used as an estimate of age. Outliers were discarded
as reading errors.
Proc. Linn. Soc. N.S.W., 130, 2009
Currently, examination of tooth structure is
the most precise method of age determination in
pinnipeds (McCann, 1993), including South African
fur seals (Oosthuizen, 1997; Oosthuizen and Bester,
1997). However, this method can only be used in
South African fur seals < 12 y. At about 12 y of age,
closure of the pulp cavity terminates tooth growth
and no further growth rings are formed. Arnould and
Warneke (2002) claim that growth rings could be
distinguished in male Australian fur seals up to 16 y
and a similar upper limit of about 15 y was found in
the Antarctic fur seal by Payne (1978). Payne (1978,
1979) also found that useful ages could be estimated
from growth lines in the cementum of the teeth of
Antarctic fur seals (A. gazella) but this method was
not attempted in the present study.
Ofthe 149 animals in the study: (1) 39 were known-
age MCM animals; (11) 34 were aged from counts of
incremental lines observed in the dentine of upper
canines, i.e., range |—11 y; (111) 10 were identified as
adults = 12 y (pulp cavity of the upper canine closed);
(iv) 66 were not aged but could be classified into
subadults and adults based upon SBL; allowing for
(1), (11) and the problem animals mentioned in (iii)
above, there was a total of 73 animals of known age
available for modelling of growth vs. age.
For this study, the following age groups were
used: pup (< | month to 6 months); yearling (7
months to 1 y 6 months); subadult (1 y 7 months to
7 y 6 months); and adult (= 7 y 7 months) (Table 1).
Very old animals of known-age were not available for
examination. Estimated longevity is c. 20 y, based
upon the lifespan of zoo animals and known life-spans
of other fur seals (Wickens, 1993). Australian male fur
seals (A. pusillus doriferus) have a lifespan of about 20
years but female Australian fur seals are known to live
to over 20 y (Arnould and Warneke, 2002). The New
Zealand fur seal (A. forsteri) (McKenzie et al., 2007)
and the Steller sea lion (Eumetopias jubatus) (Winship
et al., 2001) both have similar lifespans of about 20 y
for males and well in excess of 20 y for females.
Australian Material
The South African fur seal data on SBL vs. age
were compared to published material from Arnould
and Warneke (2002) on Australian fur seals. Data
were read off the graphs in their published paper
(Arnould and Warneke, 2002) with an accuracy of the
SBL readings of about + | cm. Fits of their data were
then compared to similar data for South African fur
seals from the present study using the same statistical
software.
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' i | |
\ | 4 MA |
} Pee ee
tte
tf \ | ee
\ | iN hey’
Biblicous rd /
\ /
\ I /
\ | . /
| |
Opening \ ¢ fe ee
\
\
\ I
\ I
;
Anus \ I
{—- —— cca Stat gt and beg |
Tail } —_ . ym en wen es
JA evel
Veo
/ Vea
/ \ |
/ \
/ \ \ 1
Lo \ Al
Figure 1: Diagram of a male South African Fur Seal showing how individual body measurements were
taken. All measurements were taken with the animal lying on its back.
B1, Circumference of head at canine; B2, circumference of head at eye; B3, tip of snout to centre
of eye; B4, tip of snout to centre of ear; B5, tip of snout to angle of gape; B6, standard body length or SBL
(straight line from tip of snout to tip of tail with animal lying on its back); B7, ventral curvilinear length
(tip of snout to tip of tail over body curve); B8, tip of snout to genital opening; B9, tip of snout to anterior
insertion of the foreflipper; B10, length of foreflipper (anterior insertion to tip of first claw); B11, axillary
girth; and B12, length of hind flipper (anterior insertion to tip of first claw). All body measurements were
made in cm.
DU) Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
Table 1: The age distribution of Male South Af-
rican Fur Seals. Pups were defined as animals <
1 month old. Animals 1-10 y: 37 MCM animals
were of known-age; 34 PEM animals were aged
from counts of incremental lines observed in the
dentine of upper canines. Animals > 12 y: 2 MCM
animals were 13 y; 10 PEM males were > 12 y, i.e.,
the pulp cavity of the upper canine was closed.
Statistical analysis
Body variable expressed in relation to standard body
length
Growth in body measurement, relative to standard
body length (SBL), was calculated as follows:
body measurement (cm)/SBL (cm) x 100%
As the variance of the ratio estimate is difficult
to validly estimate, particularly on small samples,
percentages must be interpreted with caution, i.e.,
both y and x vary from sample to sample (Cochran,
197 Te pals3):
Body length as an indicator of age
The degree of linear relationship between log
body measurement (log SBL) and age (y) was
Proc. Linn. Soc. N.S.W., 130, 2009
calculated using the Spearman rank-order correlation
coefficient.
Linear discriminant analysis can be used to
classify individual seals into mutually exclusive age
groups based on seal body length. The dependent
variable (y) is the age group and the independent
variable seal body length (x) is the feature that might
describe the age group. For each age group we can
determine the mean of seal body length (x, ) and for
each seal we compute the Mahalanobis distances of
the body length (x) to the mean seal body length of
age group 7:
EQUATION 1
Di(x)=—2|x, S"x=4%, 87x, |+ x7S x
where, S is the pooled sample variance matrix. Since
we are dealing with univariate data we have xe =X;
, X' = X and S being the pooled sample covariance.
The term in square brackets is the linear discriminant
function. We allocate an observation (x) to the age
group (pup, yearling, sub adult, adult), which gives
the smallest calculated Mahalanobis distance. This
is equivalent to allocating the observation (x) to the
age group which has the largest linear discriminant
function value (Anderson, 1984).
Growth Models
The most commonly used growth models (SBL
vs. age) for post-natal growth of marine mammals
are the exponential saturation curve, known as the
von Bertalanffy model, the logistic ‘curve and the
double exponential or Gompertz model (Zullinger et
al., 1984; Trites and Bigg, 1992, 1996; Zeide, 1993;
Winship et al., 2001; Arnould and Warneke, 2002;
McKenzie et al., 2007). In most cases where these
equations have been used, a time base adjustment
(moving the x-axis) has been used to optimise the
fit but this is not a good statistical procedure. No
attempt is usually made to estimate the errors of the
fitted parameters. In the present study, the models
have been expressed in forms where the unknowns
were the asymptotic maximum size, the apparent pup
size (P) and an exponential constant. Models are for
post-natal growth; they are not intended to model the
growth of suckling pups and the apparent pup size (P)
does not necessarily reflect the actual birth size:
223
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
EXPONENTIAL SATURATION OR VON
BERTALANFFY CURVE
EQUATION 2
Y=(E, -P)(1-e")+P
or Y=E,, -E,e™+Pe™
where, E isthe asymptotic maximum size,
Pis the apparent pup size,
kis a exponental growth constant
tis tim e.
LOGISTIC EQUATION
EQUATION 3
E
=
! iE ‘
| 1+ | Serie.
4, M, I / F
!
where,E isthe asymptotic maximum size,
é %
E,, oe aes te eee cree
| a 1 | is a scaling constant,
i
Pis the apparent pup size,
kis an exponental constant,
tis time.
DOUBLE EXPONENTIAL OR GOMPERTZ
EQUATION - EQUATION 4
a er |
wa E... = [xP )- uF Cie
where, E isthe asymptotic maximum aze,
[Ln(P)-LnfE, jlisa scaling constant
Pis the apparent pup size,
kis an exponential constant,
tis time.
For Equations 2, 3 and 4 the incremental component
of growth (sat) 1S;
EQUATION 5
gon Ms = ge
The approxim ate error for E ners (AE pew his:
, Ti SH Races
AE aonth = “f (anlet jar (AP }
where, dh
APisthe error of the apparent pup size.
224
13 the error of the maximum body size,
The growth of suckling pups would be expected
to be governed by a different growth curve and so
the apparent pup size (P) is an abstraction. There are
also statistical limitations of the models. Three (3)
unknowns have to be fitted. It is much more difficult
to fit an equation with 3 unknowns than one with 2
unknowns. The characteristics of the underlying
function can also give rise to difficulties; the logistic
equation, in particular, is notoriously difficult to fit
(Zullinger et al., 1984). The equations cannot be
adequately fitted if there is an insufficient amount
of data to clearly indicate curvature towards an
asymptotic maximum.
The errors of the fitted parameters can be
estimated using matrix inversion methods (Johnson
and Faunt, 1992). However, most attempts to use
such growth curves on mammals and growth of trees
have not used enough data points, resulting in the
asymptotic errors being so large that the estimates of
the fitted parameters are not useful (Zullinger et al.,
1984; Zeide, 1993).
Most previous attempts to fit various types
of exponential saturation curves have used data
where the equations have been simplified by using
a fixed estimate of the initial condition at t = 0 (the
apparent pup size), hence simplifying the equations
to equations with only two unknowns (Australian fur
seals - Arnould and Warneke, 2002; New Zealand
fur seals — McKenzie et al., 2007; Steller sea lion
— Winship et al., 2001).
Least squares fitting routines assume that
the error in the dependent variable is normally
distributed and independent of the magnitude of the
independent variable. In many biological situations
this assumption is not valid because the error of
the dependent variable increases with increasing
magnitude of the independent variable. A constant
relative error is often a more realistic assumption
to make for biological data. The usual procedure to
deal with situations is to log/log transform the data
and then use a least squares fitting procedure on the
transformed data. In the present study, we found
no great improvement in the curve fits (in terms of
correlation r) using log/log transformed data. Plots
of residuals vs. predicted Y-values did not indicate
a systematic increase in the size of the residuals as
the predicted Y-value increased. No log/log
transform was needed.
Bivariate allometric regression
The relationship between value of body
measurement and: (i) SBL and (11) age (y),
was investigated using the logarithmic (base e)
transformation of the allometric equation,
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
y = ax’, which may equivalently be written as log
y = log a+ b log x. ‘Robust’ regression (Huber M-
Regression) was used to fit straight lines to the
transformed data. The degree of linear relationship
between the variables was calculated using the
Spearman rank-order correlation coefficient, r
(Gibbons and Chakraborti, 1992). This is a non-
parametric procedure. Since the log-transformation is
monotonic you get the same value for r on transformed
or untransformed data. It is important to note that the
regression equations relating to overall growth are
not used on body measurements that are likely to vary
with seasonal variations in body condition that are
known to occur in this species (e.g. Rand, 1956). For
example, body girth or weight would be inappropriate
parameters to use in such analyses.
Statistical tests of hypotheses about model
parameters are only validifthe model assumptions hold
(1.e., errors are independently and identically normally
distributed, with zero mean and with a variance (o”)
(Weisberg, 1985, p. 24, 156). The standard approach
is to first examine the residual values versus fitted
plot. If this is a random scatter about zero then it is
valid to assume the model is adequate and proceed to
check the normality assumption. In the present study,
the following tests for checking for normality were
used: (i) Anderson-Darling, (ii) Ryan-Joiner and (111)
Kolmogorov-Smimov (Cochran, 1977).
We used the following test statistic to test one of
the hypotheses given below about the slopes of the
fitted lines:
EQUATION 6
—
_ b=)
SE(b)
where, bis our estimate of the slope using robust
0 regression and SE ( b ) is the standard error
of b . Under the null hypothesis the test statistic
T has a ¢ distribution with - 2 degrees of freedom
(df).
The following hypotheses were tested:
H,: b= 1 (isometric) versus H,: b # | (either positively
or negatively allometric); H,: b > 1 (positively
allometric); H,: b < 1 (negatively allometric).
Statistical Software
Statistical analysis and graphics were
implemented in Minitab (Minitab Inc., State College,
Proc. Linn. Soc. N.S.W., 130, 2009
1999, 12.23), Microsoft Excel 97 (Microsoft Corp.,
Seattle, 1997) and S-PLUS (MathSoft, Inc., Seattle,
1999, 5.1). The EXCEL 97 routines for non-linear
least squares fits and calculation of the asymptotic
errors of the fitted parameters for the von Bertalanffy,
Logistic and Gompertz equations (Equations 2, 3 and
4) are available from Dr R.J. Ritchie (rrit3 143 @usyd.
edu.au) upon request.
Terminology
A juvenile is a weaned pup that has not yet
achieved adult size. Puberty is when reproduction first
becomes possible (production of sperm in quantity),
and social maturity is the age when the animal
reaches full reproductive capacity (physically able to
establish and maintain a harem). Sexual development
of male South African fur seals is discussed elsewhere
(Stewardson et al., 1998).
RESULTS
Age determination based on dentition (intra-
observer variability)
Counts of GLGs (growth layer groups) in canine
teeth were found to be highly reproducible. Of the
34 PEM animals for which GLGs were counted, 14
(41%) had all five readings equal; 16 (47%) had one
reading out of 5 different from the mode; and 4 (12%)
had 2 readings out of 5 different from the mode.
Age determination (variability between known-
age and canine aged animals)
Standard body length (SBL) was selected to
investigate whether MCM (animals of known-age)
and PEM (canine aged animals) animals were similar
with respect to age. When comparing the (robust)
regression line for SBL on age for MCM animals with
SBL on age for PEM animals, partial t-tests indicate
that age is important (t = 7.07, p < 0.001), even after
adjusting for group and age-group interaction; but
they provide little information on group (f= -0.82, p
= 0.42) and age group interaction (¢ = 0.87, p = 0.58),
hence one straight line can be fitted to the data. These
statistical conclusions were verified by examining
graphical displays of fitted values and residuals.
Thus PEM and MCM animals were not significantly
different with respect to age distribution.
This conclusion is supported by the sequential F
test, provided the sequence of terms added sequentially
(first to last) was: (i) none (i.e., fitting a line parallel
to the x axis); (ii) age (F = 817.69, p < 0.001) (one
straight line); (iii) museum (1.e., MCM and PEM) (F
= (0.0659, p = 0.7984) (two parallel lines); (iv) age x
225
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
museum interaction (F = 0.1883, p = 0.6661) (two
lines not necessarily parallel).
Bivariate allometric regression
Regression statistics for body measurements on
SBL and age (1-10 y) are given in Appendix 3 and
4. Overall, correlation coefficients were moderately
to strongly positive, i.e., most points on the scatter
plot approximated a straight line with positive
slope, r => 0.70. Exceptions included tip of snout to
centre of eye (B3) with age and SBL (r = -0.008
and r = 0.15 respectively); tip of snout to angle of
gape (BS) with age (r = 0.56); circumference of
head at canine (B1) with age (r = 0.59). Although
correlation coefficients indicate that linearity was
reasonably well approximated for most variables
by log-log transformations, a linear relationship did
not necessarily best describe the relationship. In the
present study, we have attempted to fit more complex
models in the case of SBL vs. age with the specific
aim of comparing our growth curves with those found
for the Australian fur seal (Arnould and Warneke,
2002)(see below).
Growth of body variables
Most variables were significantly positively
correlated with each other, r = 0.68 (Appendix 2).
Exceptions were: (1) tip of snout to centre of eye (B3)
with all variables; (i1) circumference of head at eye
(B2) with tip of snout to angle of gape (B5) (r= 0.61);
and (111) circumference of head at canine (B1) with tip
of snout to angle of gape (BS5) (7 = 0.63).
Circumference of head at canine (B1)
Growth of circumference of head at canine (B1)
was variable relative to age, r = 0.59 (Appendix 4).
Overall growth expressed negative allometry relative
to SBL and age (Appendix 3, 4), increasing by 57%
at 10 y relative to pups (RTP) (Table 2). Growth
increment decreased with increasing SBL until about
7 y (c. 15% of SBL) (Table 3). The mean B1 of males
> 10 y (including unaged animals > 200 cm and of
indeterminate age > 12 y) was 31.8 + 1.2 cm (n=S).
The maximum-recorded value was 35.0 cm (animal
MCM3017, SBL 209 cm, 12 y 11 months).
Circumference of head at eye (B2)
Growth of circumference of head at eye (B2) was
rapid during the early postnatal period and continued
to increase until at least 13 y. Overall growth
expressed negative allometry relative to SBL and
scaled with negative slope relative to age (6 = 0.12)
(Figures 2a, b; Appendix 3, 4), increasing by 65% at
10 y (RTP) (Table 3). Growth increment decreased
with increasing SBL until about 7 y (c. 22% of SBL)
(Table 2). Mean B2 of males > 10 y (including unaged
animals > 200 cm and of indeterminate age > 12 y)
was 45.8 + 1.8 cm (n = 6). Maximum recorded value
was 53.0 cm (animal PEM676, SBL 197 cm).
Tip of snout to centre of eye (B3)
Growth of tip of snout to centre of eye (B3) was
highly variable relative to age, r= -0.008, and SBL, r=
0.15 (Appendix 3, 4). Growth increment decreased
with increasing SBL until about 9 y (c. 5% of SBL)
(Table 2). Mean B3 of all males > 10 y (including
unaged animals > 200 cm and of indeterminate age >
12 y) was 10.4 + 0.6 cm (7 = 10). Maximum recorded
value was 14.4 cm (animal PEM2194, SBL 194 cm).
Table 2 (Pages 227-228): Summary statistics for body variables (B1—B12), according to age (y) and age
group of male South African Fur seals.
Data presented as mean body measurement in cm + S.E., followed by coefficient of variation in
round brackets, and body variable expressed as a percentage of SBL. Maximum value of each variable
(males of unknown-age) is also presented.
Variables: B1, Circumference of head at canine; B2, circumference of head at eye; B3, tip of snout to
centre of eye; B4, tip of snout to centre of ear; B5, tip of snout to angle of gape; B6, standard body length
(SBL); B7, ventral curvilinear length; B8, tip of snout to genital opening; B9, tip of snout to anterior
insertion of the foreflipper; B10, length of foreflipper; B11, axillary girth; B12, length of hind flipper.
Variable B3 was poorly correlated with body variables and age (Appendices 1, 2, 3 and 4), therefore has
been excluded from further analysis. B7 was shown to be a poor indicator of SBL, therefore was exclud-
ed from further analysis. B11 may be influenced by seasonal change and illness, therefore was excluded
from further analysis. Sample size (n) is the number of dentition-aged and known-age (tagged) animals.
Sample size given in square brackets where this does not equal total sample size. The data summary
includes calculations of the mean of each variable + S.E. for the 7 largest males (> 200 cm) of known or
unknown-age; maximum value in square brackets, followed by sample size.
226 Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
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Proc. Linn. Soc. N.S.W., 130, 2009
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
Table 2 continued
Age group
Pup
Yearling
Subadult
Adult
Total
Mean for males >
200 cm
[max. value in
brackets]
Age (y)
=i
10
13
31.3 +2.0
[35.0] n=3
Sample size
(n)
3
10
11
45
17
73
44.34+3.2
[50.0] n=3
B7
70.9 + 3.6
(8.7) —
95.3 + 3.9 [3]
(7.1) -
149.7 +2.7 [3]
@2Q)=
155.3 +5.2 [3]
Gy
158.5 + 4.3 [5]
(6.0) -
155.2+2.6 [11]
6S)=
166.0 + 2.1 [5]
Qs3)=
185.8 + 3.4 [4]
Cu
203.7 + 4.9 [3]
(4.2) -
— [0]
182.0 + 4.9 [12]
(9.2) -
29
91.0 + 3.4
[98.0] n=4
B8
55.6 + 1.7
(5.3) 80.2%
75.9 +2.2
(8.1) 83.7%
79.6 + 2.4
(6.8) 84.9%
98.1+42.1
(4.9) 87.0%
107.2 + 3.6 [8]
(9.5) 86.2%
124.5+4.8
(8.7) 84.4%
WGA 22 M7)
(4.2) 87.2%
3-5 2e 25)
(6.3) 84.9%
115.7 + 2.9 [44]
(16.5) 86.0%
136.6 + 3.1
(5.5) 85.8%
152.6+ 2.6
(3.8) 89.6%
159.3+5.4
(6.8) 87.1%
178.5 + 3.5
(2.8) 86.4%
151.6 + 3.8
(10.2) 87.4%
72
49.0 + 2.7
[55.0] n=4
B9
31.7+0.9
(4.8) 45.7%
Al = 1.7
(11.6) 45.3%
37.7+£0.8
(4.6) 40.2%
48.9+2.1
(9.5) 43.3%
52.5 + 1.7
(9.8) 42.6%
62.7442
(14.9) 40.5%
65.5 + 3.4
(16.3) 43.6%
71.8+2.1
(9.6) 45.8%
59.2+2.0
(22.2) 43.4%
76.6 + 2.2
(7.1) 50.1%
83.8+5.8
(15.6) 48.4%
87.8 + 8.5
(19:5) 48.1%
91.5+3.5
(5.4) 44.3%
83.142.9
(14.1) 47.9%
73
135.0 + 34.0
[169.0] n=2
B10
17.6+ 1.6
(16.2) 25.4%
22.4+ 1.2
(15.0) 24.7%
23.5+0.4
(4.3) 25.1%
27.4+£1.4
(11.2) 24.3%
30.1+ 1.1
(11.0) 24.5%
35). 7/ 2 IA!
(9.0) 23.9%
33.6 + 0.9 [9]
(7.7) 23.0%
34.7+ 1.1
(10.1) 22.4%
31.6 + 0.7 [44]
(15.3) 23.6%
35.2+1.6
(11.5) 21.3%
40.6 + 0.9
(5.0) 24.1%
40.3 + 2.1 [3]
(10.2) 22.0%
48.4 + 3.6
(10.5) 23.4%
39.5 + 1.3 [16]
(13.5) 22.7%
72
28.8 + 1.4
[29.2] n=3
Bll
39.6 +3.5
(15.5) 57.1%
53.14£4.6
(24.4) 58.5%
58.2431
(11.8) 62.0%
73.9 +2.0
(6.2) 65.5%
80.2+2.3
(8.5) 64.7%
85.8 + 0.8 [2]
(1.2) 62.8%
91.4+2.1 [9]
(6.8) 62.7%
100.2 + 3.1 [7]
(8.3) 64.4%
83.2+2.4 [37]
(17.5) 63.8%
90.6 + 4.6 [2]
(7.2) 57.5%
114.5 +2.9 [4]
(5.1) 67.0%
111.9+6.9
(12.2) 61.2%
— [0]
108.7+4.1 [10]
(12.0) 62.8%
58
B12
13.3£0.7
(9.4) 19.2%
15.1+£0.4
(8.0) 16.6%
16.0 + 0.6
(8.3) 17.0%
18.1408
(9.3) 16.0%
18.6 + 0.5
(8.8) 15.2%
21.7+1.3 [3]
(10.1) 15.0%
21.2 +0.4 [9]
(5.9) 14.5%
23.6 +0.7 [10]
(9.1) 15.2%
20.2 + 0.5 [41]
(15.0) 15.3%
26.0 + 1.0
(9.7) 15.8%
28.2413
(10.1) 16.5%
27.1 + 1.6
(11.7) 14.8%
27.7£1.5
(7.7) 13.4%
27.1 £0.46
(9.8) 15.3%
Proc. Linn. Soc. N.S.W., 130, 2009
69
228
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
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Proc. Linn. Soc. N.S.W., 130, 2009
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
Tip of snout to centre of ear (B4)
Growth of tip of snout to centre of ear (B4) was
rapid during the early postnatal period and continued
to increase until at least 13 y (Table 2 and 3). Overall
growth expressed negative allometry relative to
SBL and scaled with negative slope relative to age
(b = 0.04) (Figures 3a, b; Appendix 3, 4), increasing
by 70% at 10 y RTP (Table 3). Growth increment
decreased with increasing SBL until about 7 y (c.
12% of SBL) (Table 2). The mean B4 of all males
> 10 y (including unaged animals > 200 cm and of
indeterminate age > 12 y) was 22.7 + 0.8 cm (n = 7).
The maximum-recorded value was 25.2 cm (animal
MCM3125, SBL 204 cm, 13 y).
Tip of snout to angle of gape (B5)
Growth of tip of snout to angle of gape (B5)
was variable relative to age, r = 0.56 (Appendix 4).
Overall growth scaled with negative slope relative
to SBL (6 = 0.64) and expressed negative allometry
relative to age (Appendix 3, 4), increasing by 55%
at 10 y RTP (Table 3). Growth increment decreased
with increasing SBL until about 7 y (c. 6% of SBL)
(Table 2). The mean BS5 of all males > 10 y (including
unaged animals > 200 cm and of indeterminate age
> 12 y) was 13.2 + 0.7 cm (n = 7). The maximum
recorded value was 15.0 cm (animal PEM676, SBL
ISS eran),
Standard body length (B6 or SBL)
Growth of SBL (B6) was rapid during the early
postnatal period with a significant growth spurt
between 2 and 3 y (two sample t test: p-value = 0.008;
df = 5). The rate of growth slowed significantly
between 6 and 7 y (two sample t test assuming
unequal variances: p-value = 0.011; df= 9). A weak
growth spurt was observed at 9 and 10 y but could
not be examined statistically, 1.e., this secondary
growth spurt may be attributed to sampling error.
Growth increased by 164% at 10 y RTP (Table 3).
Considering that the 13 y old males measured 206.5
+ 2.5 cm (n = 2), and mean SBL of all males > 10 y
and/or unaged animals > 200 cm was 197 = 4.1 cm
(n = 15), growth appears to slow after attainment of
social maturity (Table 2).
Tip of snout to genital opening (B8)
Growth of tip of snout to genital opening (B8)
was rapid during the early postnatal period and
continued to increase until at least 13 y (Table 2 and
3). Growth increased by 186% at 10 y RTP (Table
3). In subadults and adults, mean value remained at
about 86% of SBL (Table 2). Overall growth scaled
with weak positive slope relative to SBL (6 = 1.04)
and negative slope relative to age (6 = 0.02). The
maximum recorded value for parameter B8 was 184.0
cm (animal PEM2256, SBL 198 cm). The mean B8
of all males > 10 y, including unaged animals > 200
cm) was 171.143.4cm (n=7).
Tip of snout to anterior insertion of the
foreflipper (B9)
Growth of tip of snout to anterior insertion of the
foreflipper (B9) was rapid during the early postnatal
period and continued to increase until at least 10 y
(Table 2 and 3). Overall growth expressed positive
allometry relative to SBL, and negative allometry
relative to age (Figure 4a, b; Appendix 3, 4). Growth
increased by 177% at 10 y RTP (Table 3). Mean SBL
of all males > 10 y, including unaged animals > 200
cm was 94.2 + 3.1 cm (m = 7). Maximum recorded
value for B9 was 110.0 cm (animal PEM2374, SBL
186 cm).
Length of foreflipper (B10)
Growth of length of foreflipper (B10) was rapid
during the early postnatal period and continued to
increase until at least 13 y (Table 2 and 3). A significant
growth increment was evident between 4 and 5 y (two
sample t test: p-value = 0.015; df= 8). Overall growth
scaled with negative slope relative to SBL (b = 0.89)
and age (b = 0.07). Growth increased by 129% at 10
y RTP (Table 3). Growth increment decreased with
increasing SBL until about 6 y (c. 23% of SBL) (Table
2). The mean length of flipper (B10) of all males > 10
y, including unaged animals > 200 cm was 47.2 + 1.9
Figure 2a, b (right): Bivariate plot of log circumference of head at canine (cm) on: (a) log SBL length
of seal (cm) and (b) age (y). PEM animals, open squares; MSM animals, closed triangles.
Figure 3a, b (right: Bivariate plot of log tip of snout to centre of ear (cm) on: (a) log length of seal (cm)
and (b) age (y). PEM animals, open squares; MCM animals, closed triangles.
Figure 4a, b (right): Bivariate plot of log tip of snout to anterior insertion of the foreflipper (cm) on: (a)
log length of seal (cm) and (b) age (y). PEM animals, open squares; MCM animals, closed triangles.
230
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
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