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.
The current subscription rate to the Proceedings is set at A$110.00 per volume. In recent years
a volume consists of a single annual issue.
Back issues of all but a few volumes and parts of the Proceedings are available for purchase. Prices are
listed on our home page and can also be obtained from the Secretary.
OFFICERS AND COUNCIL 2007/2008
President: Michele Cotton
Vice-presidents: M.L. Augee, I.G. Percival, D.R. Murray
Treasurer: 1.G. Percival
Secretary: J-C. Herremans
Council: A.E.J. Andrews, M.L. Augee, J.P. Barkas, M. Cotton, M.R. Gray, J-Cl. Herremans, D. Keith,
R.J. King, H.A. Martin, E. May, D.R. Murray, Joelle Myerscough, I.G. Percival, J. Pickett, S. Rose, H.M.
Smith and K.L. Wilson
Editor: M.L. Augee
Assistant Editor: Elizabeth May
Auditors: Phil Williams Carbonara
The postal address of the Society is: P.O. 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.
Dumeril, A.M.C., Bibron, G. and Dumeril, A. (1854).
Erpétologie Générale ou Histoire Naturelle Complete
des Reptiles. Paris: Roret Vol. 7(2) pp. 781-1536 [p.
1254-1258, pl. 76 b. fig. 1].
Fletcher, J.J. (1929). The Society’s heritage from the
Macleays. Proceedings of the Linnean Society of New
South Wales 54, 185-272. [269]
Goldingay, R.L. (1998). Between a rock and a hard place:
conserving the broad-headed snake in Australia’s
oldest National Park. Proceedings of the Linnean
Society of New South Wales 120, 1-10.
Goldingay, R.L. and Newell, D.A. (2000). Experimental
rock outcrops reveal continuing habitat disturbance
for an endangered Australian snake. Conservation
Biology 14, 1908-1912. ~
Guibé, J. and Roux-Estéve, R. (1972). Les types de
Schlegel (Ophidiens) présents dans les collections
du Muséum National d’ Histoire Naturelle de Paris.
Zoologische Mededeelingen 47: 129-134.
Giinther, A. (1858). Catalogue of colubrine snakes in the
collection of the British Museum. London: British
Museum. Pp. 213-214.
Hersey, F. (1980). Broad-headed snake Hoplocephalus
bungaroides. Pp. 38-40, in Endangered Animals of
New South Wales, edited by C. Haigh. National Parks
and Wildlife Service, Sydney.
Hoser, R. (1995). The Australian broad-headed snake
(Hoplocephalus bungaroides). The Reptilian
Magazine 3(10), 15-27.
Keogh, J.S. (1998). Molecular phylogeny of elapid snakes
and a consideration of their biogeographic history.
Biological Journal of the Linnean Society 63, 177-
203.
Keogh, J.S. (1999). Evolutionary implications of
hemipenial morphology in the terrestrial Australian
MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE
elapid snakes. Zoological Journal of the Linnean
Society 125, 239-278.
Krefft, G. (1866). On snakes observed in the
neighbourhood of Sydney. Transactions of the
Philosophical Society of New South Wales 1862-
1865, 34-60.
Krefft, G. (1869). Broad-headed snake. In ‘The snakes of
Australia; an illustrated and descriptive catalogue of
all the known species’. Government Printer: Sydney.
pp. 56-57 and Plate VI, figs 6, 6a, 6b.
Leviton, A.E., McDiarmid, R., Moody, S., Nickerson, M.,
Rosado, J., Sokal, O., and Voris, H. (1980). Museum
acronyms — second edition. Herpetological Review
11, 93-102.
Leviton, A.E., Gibbs, R.H. Jr., Heal, E., and Dawson, C.E.
(1985). Standards in herpetology and ichthyology:
Part I. Standard symbolic codes for institutional
resource collections in herpetology and ichthyology.
Copeia 1985(3), 802-832.
Longmore, R. (1989). Hoplocephalus bungaroides.
In ‘Snakes: atlas of elapid snakes of Australia.’
Australian Flora and Fauna Series No. 7. Australian
Government Printing Service, Canberra. p. 64.
Loveridge, A. (1934). Australian reptiles in the Museum
of Comparative Zoology Cambridge Mass. Bulletin
of the Museum of Comparative Zoology (Harvard
College) 77(6): 243-383. [290]
Newell, D.A. and Goldingay, R.L. (2005). Distribution
and habitat assessment of the broad-headed snake
(Hoplocephalus bungaroides). Australian Zoologist
33, 168-179.
Reiling, H. and Spunarova, T. (2005). Vaclav Fri¢ (1839—
1916) and his influence on collecting natural history.
Journal of the History of Collections 17, 23-43.
Roselaar, C.S. (2003). An inventory of major European
bird collections. Bulletin of the British Ornithologists’
Club 123A, 253-337.
Schlegel, H. (1837). Essai sur la Physionomie des
Serpens. Partie Générale et Partie Descriptive. La
Haye: Kips and Stockum 2, 477-478.
Shaffer, H.B., Fisher, R.N., and Davidson, C. (1998). The
role of natural history collections in documenting
species declines. Trends in Ecology and Evolution 13,
27-30.
Shea, G.M. and Sadlier, R.A. (1999). A catalogue of the
non-fossil amphibian and reptile type specimens
in the collection of the Australian Museum: types
currently, previously and purportedly present.
Technical Reports of the Australian Museum 15, 1-
Dil.
Shine, R. (1981a). Venomous snakes in cold climates:
ecology of the Australian genus Drysdalia
(Serpentes: Elapidae). Copeia 1981, 14-25.
Shine, R. (1981b). Ecology of the Australian elapid snakes
of the genera Furina and Glyphodon. Journal of
Herpetology 15, 219-224.
Shine, R. (1983). Arboreality in snakes: ecology of the
Australian elapid genus Hoplocephalus. Copeia
1983, 198-205.
10
Shine, R. and Fitzgerald, M. (1989). Conservation
and reproduction of an endangered species: the
broad-headed snake, Hoplocephalus bungaroides
(Elapidae). Australian Zoologist 25, 65-67.
Shine, R., Webb, J.K., Fitzgerald, M. and Sumner,
J. (1998). The impact of bush-rock removal on
an endangered-snake species, Hoplocephalus
bungaroides (Serpentes: Elapidae). Wildlife Research
25, 285-295.
Slowinski, J.B. and Keogh, J.S. (2000). Phylogenetic
relationships of elapid snakes based on cytochrome
b mtDNA sequences. Molecular Phylogenetics and
Evolution 15, 157-164.
Strahan, R. (1979). Rare and curious specimens: an
illustrated history of the Australian Museum 1827-
1979. The Australian Museum Trust.
Swan, G., Shea, G. and Sadlier, R. (2004). A field guide to
reptiles of New South Wales. 2nd Edition. Reed New
Holland.
Wallach, V. 1985. A cladistic analysis of the terrestrial
Australian Elapidae, pp. 223—253 in Biology of
Australasian frogs and reptiles (G. Grigg, R. Shine,
and H. Ehmann, eds.). Royal Zoological Society of
New South Wales, Chipping Norton.
Wallach, V. 1998. The lungs of snakes, pp. 93-295 in
Biology of the Reptilia (C. Gans and A. S. Gaunt,
eds.), Vol. 19 (Morphology G). Society for the Study
of Amphibians and Reptiles, Athens.
Webb, J.K. and Shine, R. (1997). Out on a limb:
conservation implications of tree-hollow use
by a threatened snake species (Hoplocephalus
bungaroides: Serpentes, Elapidae). Biological
Conservation 81, 21-33.
Webb, J.K. and Shine, R. (1998a). Using thermal ecology
to predict retreat-site selection by an endangered
snake species (Hoplocephalus bungaroides:
Serpentes, Elapidae). Biological Conservation 86,
233-42.
Webb, J.K. and Shine, R. (1998b). Ecological
characteristics of a threatened snake species,
Hoplocephalus bungaroides (Serpentes, Elapidae).
Animal Conservation 1, 185-93.
Webb, J.K. and Shine, R. (2000). Paving the way for
habitat restoration: can artificial rocks restore
degraded habitats of endangered reptiles? Biological
Conservation 92, 93-99.
Webb, J.K., Brook, B.W. and Shine, R. (2002). Reptile
collectors threaten Australia’s most endangered
snake, the broad-headed snake Hoplocephalus
bungaroides. Oryx 36, 170-181.
Whitley, G.P. (1961). The life and work of Gerard Krefft.
Proceedings of the Royal Zoological Society of New
South Wales 1958-59, 21-34.
Whitley, G.P. (1969). Gerard Krefft (1830-1881) and his
bibliography. Proceedings of the Royal Zoological
Society of New South Wales 1967-68, 38-42.
Proc. Linn. Soc. N.S.W., 130, 2009
J.M.HARRIS AND R.L. GOLDINGAY
yno juas
sordeds jo sisi] ou ‘AmnqioyUR_ ‘SUIT Il WOT, POAlooel sem (Saploimsung T=) ..Snjoselipa srypydaosojdopy ayxeuUs Yoe[q poyesoisea V,,
“SOSULYOXO JO SI] Be SUIdsoy UdEq jou A[O}e]
pey yFory yeu odes jenuue sty} Ul poyeys SoojsnI] JO pleog oy, popraoid a19M sasueLYyOXe JO S}sI{ OU PUR ‘poATodel o19M SaplOADSUNG "FT ON
“SniyLineyy “SNOT LO “Pse][IGOY op ‘A Il 0} JUSS se
sander 67 JO WONeT[09 V ‘Ad|PeIg “H'H IW Wop poataoel sem (sjosalspA “YY Se) Saploapsung “FF SUIPN{OUL SoxeUS SUIAT] JO UONIeT[00 WV
‘(SNJDBALADA “FT Se) Saplospsung ‘FT 210 10 9UO SUIpNyoUT
« Ul[iog Ul unasnyy [eAOY SY} FO 10jDeM ‘S19}9q JOSSOJOIg,, 0} JUSS a1OM SaydaI JO UOTIET]OD B Inq ‘{paATedeI BIOM Saplos‘Dsung FY ON
‘[ro[vop eB] TyNowAaM “UOTURG VOqQOY IPI 0} JUSS OOM SoYsy pue spiig ‘soyydor ‘sjeurumew Jo suowmoods
OSI pure ‘[s9[vop e] onBe1g ‘QI “A I 0} JAS (Sastoj10} 7] snjd) sender 17 Swnesny] YsHIIg 0} JUSS soysy pue sojydor Jo susuriseds gz]
‘AVISIOATUL) pleAILY ‘ASO[OO7Z sATyeredui0D Jo winesny] 0} JUAS dJaM sopdar poyloadsun ¢Z JOAOMOY ‘PpoATodeI OJOM Saplo/nsuNng FY ON
‘JNO JUSS O19M SoyIda1 OU PUR ‘PpaATIdOI 19M SaploADBUNG “ET ON
‘quas satoads ay} Jo AyUSpI JO SJoquINU sy} JO
POprAold si UOTJVoIpUT ON ‘WINESNy[ UI[JOg OU} 0} JUES SEA SOYs pue sopdar ‘s]eUUeUT JO UONIETIOD & JN {poATodeI BOM SAp1O/DSUNG FY ON
‘(eIpuy ‘TeuuayD=) seIpey “S}So10,J JO 1OJeAIOSUOD,
[elouZO ‘ewoppeg “HY urleyde_ 0} ques ATpoysodar sem (snjosaisva snpoydasojdoy se) usuttoads duo {paAtade1 d19M Saplospsung “FT ON
‘(sopydor ¢Z) uel UL UMesnyy
dIAIZ ‘(soysy. pue sopyder p¢) unesnyy sted “(soysy pue soyydoe1 Q¢) Sinquiey ‘ules “TO IP “(e[Hder suo) prupeyy ye Winesnyy [eAOY
‘(soda QZ) elueusey Jo AJa1IN0g [eAOY dy} 0} JUSM soTdoy UdAIs JOU sem soapdar asoy) JO AIMUAPr 9Y} JNG SUINOSNW SNOTLIVA O} JNO yds
d1oM sopydoy ‘soroods J1oyjOUe 10 sapIOADSUN SEA SIU} IOYJOYM Poje}s JOU SLM I JNQ “URULIOH “AA IJ] & WOIF poAtooos sem snjpoydasojdoy VW
‘(WINIS[ag Ul Jo[eap &) s[nd “Of IA & 0} pue “ummasny] sueg oy} 0} JUOS oJOM ‘saplospsung “FOU yng ‘soynde1
JOJO “GOT Ul OS[Y “Uk]IP Ul UNESNY] OIAID 0} JUAS seM (SNJDSaIIDA “FT Se) SaplosDSung ‘FT 9UO yNQ ‘PaATode1 SJOM SaplosDBUNg “FT ON
‘popnyour jou Ajuoredde sem saplospsung ‘YH Yysnoujye ‘seipeyy pure ueploy “sinquieyy Ul swWMasnUT 0} JUES o19M so[Nndoy
“suOSIeg “J QUI[OIeD ssi & Ag payeuop sem (sjHSaLIDA +7) BYVUS popeoy-peolg V
“UeUaIO_ I] & Ag poyeuop sem (s7jpsal1DA snyoydesojdopy] se) axeUus popesy-peolg V
“poysijqnd jou arom sjiejap oytoads yng wMasnyl URTTeNSNY 9} 10F (Soroads QT ‘sayeus ¢¢ SuIpNjour) sojydo1 pojoo][0o YJory
s}1odoy [wnuuy WO.y po}e]]09 sa}0N
‘saplospsung snjpydasojdozy ayeus papvosy-pvo.A IY) 0} 9dUd1EJo1 UO sIseyduo ue
gould sajou oy], ‘sosuBYyoXe ofIdo1 Jo s[ivJop YIM PLST-LORT WNesny] ULTL.Qsny sy) Jo sj1odey [enuuy oy) Wor Usye) Soj0u ArevUIUINS :] xIpueddy
pL81
eL81
CL81
IL81
OL81
6981
8981
L981
9981
£981
y98l
€981
C981
1981
TBI
I]
Proc. Linn. Soc. N.S.W., 130, 2009
MUSEUM HOLDINGS OF THE BROAD-HEADED SNAKE
BIZAIOT JO AUISIOAIUL) “WOTJIET[OD [eo1d0[007
- USpegsolM “SUNTWIUUILS oYOIPyeYyOsuossIMInjeN] USpeqso}:l - WNOISNUIS]JG Sio}s[ey-pueljoy D>
- JoJsuny “OpunyzINjeN] Inj wunasnyy] SoyosTpeyIsoM, = snyry “Winosny] SI10}sTyINye NY =
ci SloMyosuneig “WInesnyy SOYOSLIO}STYINJeN] SOYIITIeLIS yievuuseg GN
oi uapsalq “opunyxJSL], Inj winesnyy soyorperys e, onseig ‘winasny] [euoTjeN Ss
. yes}N1g ‘opunsxzINjeN Inj UNasnyy soysip}eR}S 2 ouIg “WInesny] UPIARIOT @
i oYMsj1ey “OpunyzINyeNy Iny wMesnyy soyorperis ayqndoay yoozD 5
s Z\IOH ‘opunyinjeny Inf unasnyy soyorjerys e ASO[OOZ JO winasnj] eo] Vy JO AjIsIOATU) =
- JOAOUURH “WunesnuUsapueT SayosisyoRsiapalN, os unasny] O11ej}UGE [eAOY WN
- sInqua}[y ‘WNnuURLINneyy] Wnesnyy soyorppunyinjyeNy se unesny] VIQUIN[OD Ysig [eAo yy Z,
“ SINQOD “UInesny-OpunyINje Ny e umnasnyy eyIoq] Vy [PAO
- Sizdio7] wWnesnwepunyinjeN, e daqangd “[RaNUO] ‘AVISIOATUL) [[L DOV “uMesnyy yyedpoy 5
x }pe}sjopny “WMosnyAy SoySst10}sTyIN}eN] x AIO}SIF{ [eINyeN] JO WNasnyy e109 PAON DN
. Zuley] “WINasNy] SOYOSTIO}SIYAN}e | 2, BIQUIN]OD YsHIg JO AjISIOATUL) “UANESNIA] OeIGQOLIAA\ URMOD ros
- AjeI}S|slosuosI[Ioy] ‘sIngounT NAVN be onjeN{ JO winasny] ueIpeueD &
Fs uuog “sIus0y winesnyy upeueD a
- }pe}sioq|ey Wnuvoulsy] winesnyy = BIOS “AIO\sTIH [eINJeN JO wWinesnyy [euoeNY oO
- punwyI0g ‘opunyinjeNy Iny winesnyl vlivsng °
- uazjoyuosuey ‘pjay[y winesnyy = S[OSSNIg “SsoUdIOS [Injen JO aNjYsuy wuNIsjog [eAOY Ay
- SINGSPIequ[IA\ ‘Winosny\-einp 7 USANT POISIOATUL) IYoT[ouIey “oIs0]OO7Z unosny
a }pejsueg “WinosnwIsapueT soyosIssoy] : AJISIOAIUL) JUSH “apuNns.1o1q 100A wunesny
* IVWISID “INjeN] op sneyy * S[oSsnuig ‘a1d0]007 ap winesny/y
- yoniquieT “winesnyj-rouruin qd = IVUINOL, “o]JOINJeNY OA10}STFY.P Sasnyy
Auvultasy umnisjog
- BINOQSE.NS “d1d0[00Z op dasny\[ - ULIQUIOG NVYSsInjeN] 1oS19q].1V.10/\,
c ISNO[NOT, “S][o1NjeNY O10}STEY.P UNasnyy e ZUIT “WINSsNUIsapuLT SOYOSIYdIo119}S0.10qO
x UoNoOY “o][o1njJeNY O10}STF] Pp wWinasny| 2 ZeIQ) “WINoUUeOL WINesnuUsopue Ty]
- DOIN ‘a][OINJVNY SILO}STFY.P Uunasnyy zn SINQZ|eS “Injen Jop sney
‘ SoJURN] “S][OANJVNY SILO}SIF{,.P UNasnypy BLysny
2; SO][LsIVJ] “O][OINJeNY OAO}STFY,P wunosnyj| . 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! AS ee &
wien ten
“uy |
Qh ¥%
etree, AY,
istity
Sadia
pies
ae eae
ior aT
WA Pe entered Liege aenaniey
OORT sutely tee MD
stihl Ante A
“wero ay
aes May"
iA )
“a
Ae |
yw
£#
Cd
} ‘fe ¥
byt
WE END
edinfed WAEMD
clartle® yamadl)
wha
Kyte eg
ME
MAL
rackklit Maser
#2) (OVW id lywiy Otte
i TPT mt hereon
qt rex
ia
Mia bots
rt
y
iy y
4
+e ‘
we! UME
Nea Gin {
4 ‘ Kr ry) ; ”
iy : " im LP ‘
oo) oe
a
DAMN
BMNY
hy
a bs (Ri Levaw Ap Pullen bolt sunt
eee Ne
Wetqesagid’D tian A
4 edit ingest
— oe Sa Wogan 2
Yara, Vital. sate
Viusteiens nvdue'
Pade 114)
grok 49°
ome ef “tam Neen 6. a *
dpe (ethan Abad ma ; hee, Hava, 4 Ps
Kc goto AMR
Acs ol amet “ Trew ie anes ie ‘* “
as) ae
OSTA
ve POPOL EM
ae serrate town esr REEL
md apis, aA» iy eee A hw merci a
Whewke, Vi dion =a ‘Daksa
Cay piv) : vy pet art Soil weer: tts OREO
UW Had une. pani j a petew
FEE. 64 - (20K
or Ajnctdlia,” gentnd wade
iT Yoo TiGlas orread 7 incest
ei Atel tama ety Fl lect (yur Agente eet
ai (nite mono Ear
ATK hong Baw -wanby2 iveh
yup thy Mickite ir ep Bbeh
y C0 @R5 nl Wal Mi eandeA HOSE
; Gs oGiadtiaryd baad OFRES
1 asad) Melerahury wiles s rerea
re) Com TRongalfeymamy Bay | aatme
DRS ohsbA ce bietimooed babicsA brat]
AY WaT yonbv2 TKK
Re NS cilenmanA hye RISPOR
Le BN youl? fo C1Sa0a.
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. University of New South Wales Quarterly
22, 6-8.
Archer, M. (1984). The Australian marsupial radiation.
In “Vertebrate zoogeography and evolution in
Australasia’ (Eds M. Archer and G. Clayton) pp. 633-
808. (Hesperian Press: Carlisle).
Archer, M. and Bartholomai, A. (1978). Tertiary mammals
of Australia: a synoptic view. Alcheringa 2, 1-19.
Archer, M., Hand, S. and Godthelp, H. (1995).
‘Riversleigh: the story of animals in Australia’s
ancient inland rainforests’. (Reed Books:
Chatswood).
Bakker, R.T. (1988). “The dinosaur heresies’ (Penguin
Books: London). ;
Banks, M., Colhoun, E. and Van de Geer, G. (1976).
Late Quaternary Palorchestes azael (Mammalia,
Diprotodontidae) from north-west Tasmania.
Alcheringa 1, 159-166.
Bartholomai, A. (1962). A new species of Thylacoleo and
notes on some caudal vertebrae of Palorchestes azael.
Memoirs of the Queensland Museum 14, 33-40.
Bartholomai, A. (1978). The rostrum in Palorchestes
Owen (Marsupialia: Diprotodontidae). Results of the
Ray E. Lemley expeditions. Part 3. Memoirs of the
Queensland Museum 18, 145-149.
Berganini, D. (1964). ‘The land and wildlife of Australia’.
(Life Nature Library: New York).
Black, K. (1997a). A new species of Palorchestidae
(Marsupialia) from the late middle to early late
Miocene Encore Local Fauna, Riversleigh,
northwestern Queensland. Memoirs of the
Queensland Museum 41, 181-185.
Black, K. (1997b). Diversity and biostratigraphy of the
Diprotodontoidea of Riversleigh, northwestern
Queensland. Memoirs of the Queensland Museum 41,
187-192.
33
RECONSTRUCTING PALORCHESTES
Black, K. and Mackness, B.S. (1999). Diversity and
relationships of diprotodontid marsupials Jn Archer,
M., Arena, R., Bassarova, M., Black, K., Brammall,
J., Cook, B., Creaser, P., Crosby, K., Gillespie,
A., Godthelp, H., Gott, M., Hand, S.J., Kear, B.,
Kirkman, A., Mackness, B., Muirhead, J., Musser, A.,
Myers, T., Pledge, N., Wang, Y. and Wroe, S. (Eds)
The evolutionary history and diversity of Australian
mammals. Australian Mammalogy 21, 1-45.
Buckland, W. (1824). Notice on the Megalosaurus, or
great fossil lizard of Stonesfield. Transactions of the
Geological Society 1, 390-396.
Calaby, J.H. and Lewis, D.J. (1977) The Tasmanian devil
in Arnhem Land rock art. Mankind 11, 150-151.
Chaloupka, C. (1993). “Journey in time’. (Reed Books:
Sydney).
Clegg, J. (1978). Pictures of striped animals: which
ones are thylacines? Archaeology and Physical
Anthropology in Oceania 13, 19-29.
Clegg, J. (1981). “Notes towards Mathesis art’. (Clegg
Calendars: Balmain).
Coleman, W. (1964). “Georges Cuvier: zoologist. A study
in the history of evolution’. (Harvard University
Press: Cambridge, Massachusetts).
Corrigan, P. (1988). ‘Innocent stupidities’: de-picturing
(human) nature. On hopeful resistances and possible
refusals: celebrating difference(s) —c again. In
‘Picturing power: visual depiction and social
relations’ Sociological Review Monograph 35.
(Eds G. Fyfe and J. Law) pp. 255-281. (Routledge:
London).
Delair, J.B. and Sarjeant, W.A.S. (1975). The earliest
discoveries of dinosaurs. Jsis 66, 4-25.
Desmond, A.J. (1975). ‘The hot-blooded dinosaurs. A
revolution in palaeontology’. (Blond and Briggs:
London).
Desmond, A.J. (1979). Designing the dinosaur. Jsis 70,
224-234.
Desmond, A.J. (1982). “Archetypes and ancestors.
Palaeontology in Victorian London 1850-1875’.
(Blond and Briggs: London).
De Vis, C.W. (1895). A review of the fossil jaws of
the Macropodidae in the Queensland Museum.
Proceedings of the Linnean Society of New South
Wales 10, 75-133.
Flannery, T.F. (1983). A unique trunked giant Palorchestes.
In ‘Prehistoric animals of Australia’ (Eds S. Quirk
and M. Archer) pp. 54-55. (Australian Museum:
Sydney).
Flannery, T.F. (1992). New Pleistocene marsupials
(Macropodidae, Diprotodontidae) from subalpine
habitats in Irian Jaya, Indonesia. Alcheringa 16, 321-
Boe
Flannery, T. (1994). ‘The future eaters’. (Reed Books:
Sydney).
Flannery T.F. and Archer, M. (1985). Palorchestes Owen,
1874. Large and small palorchestids. In ‘Kadimakara.
Extinct vertebrates of Australia’ (Eds P.V. Rich, G.
van Tets and F. Knight) pp. 234-239. (Pioneer Design
Studio: Lilydale).
34
Fletcher, H.O. (1945). Palorchestes - Australia’s extinct
giant kangaroo. Australian Museum Magazine 8,
361-365.
Flood, J.M. (1997). ‘Rock art of the Dreaming: Images of
ancient Australia’ (Angus andRobertson: Sydney).
Ford, J. (1982). The strange creatures of ancient Australia.
Omega Science Digest July/August, 84-87.
Fyfe, G. and Law, J. (1988). Introduction: on the
invisibility of the visual. In “Picturing power: visual
depiction and social relations’ Sociological Review
Monograph 35. (Eds G. Fyfe and J. Law) pp. 1-14.
(Routledge: London).
Gillespie, R. (2004). First and last: dating people and
extinct animals in Australia. Australian Aboriginal
Studies 1, 97-101.
Gould, S.J. (1991). “Bully for Brontosaurus’. (Penguin
Books: London).
Gregory, J.W. (1902). Some remains of an extinct
kangaroo in the dune-rock of the Sorrento Peninsula,
Victoria. Proceedings of the Royal Society of Victoria
14, 139-144.
Hall, T.S. and Pritchard, G.B. (1897). Note on a tooth of
Palorchestes from Beaumaris. Proceedings of the
Royal Society of Victoria 10, 57-59.
Hocknull, S.A., Zhao, J., Feng, Y. & Webb, G.E. (2007).
Responses of Quaternary rainforest vertebrates to
climate change. Earth and Planetary Letters 264,
317-331.
Horton, D.R. (1979). The great megafaunal extinction
debate: 1879-1979. Artefact 4, 11-25.
Horton, D.R. (1980). A review of the extinction question:
man, climate and megafauna. Archaeology and
Physical Anthropology in Oceania 15, 86-97.
Horton, D.R. (2000). “The pure state of nature: Sacred
cows, destructive myths and the environment’. (Allen
and Unwin: Sydney).
Latour, B. (1986). Visualisation and cognition. Thinking
with eyes and hands. Knowledge and society: Studies
in the sociology of culture past and present 6, \-40.
Latour, B. (1987). “Science in action. How to follow
scientists and engineers through society’. (Harvard
University Press: Cambridge, Massachusetts).
Leidy, J. (1858). Remarks concerning Hadrosaurus.
Proceeding of the National Academy of Sciences 10,
215-218.
Lewis, D.J. (1977). More striped designs in Arnhem
Land rock paintings. Archaeology and Physical
Anthropology in Oceania 12, 98-111.
Lewis, D.J. (1986). Comment ‘The Dreamtime animals’: a
reply. Archaeology in Oceania 2, 140-145.
Long, J., Archer, M., Flannery, T. and Hand, S. (2003).
‘Prehistoric mammals of Australia and New Guinea’.
(University of New South Wales Press: Sydney).
Lord, C. and Scott, H.H. (1924). “A synopsis of the
vertebrate animals of Tasmania’. (Oldham, Beddone
and Meredith: Hobart).
Lydekker, R. (1887). Catalogue of the fossil Mammalia in
the British Museum (Natural History). Part 5. (Taylor
and Francis: London).
Proc. Linn. Soc. N.S.W., 130, 2009
B.S. MACKNESS
Mackness, B.S. (1987). ‘Prehistoric Australia. 4 000
million years of evolution in Australia’. (Golden
Press: Sydney).
Mackness, B.S. (1995). Palorchestes selestiae, a new
species of palorchestid marsupial from the early
Pliocene Bluff Downs Local Fauna, northeastern
Queensland. Memoirs of the Queensland Museum 38,
603-609.
Mahoney, J.A. and Ride, W.D.L. (1975) ‘Index to the
genera and species of fossil Mammalia described
from Australian and New Guinea between 1838
and 1968’. Western Australian Museum Special
Publication 6, 1-250.
Mantell, G.A. (1825). Notice on the iguanodon, a newly
discovered fossil reptile from the sandstone of Tilgate
Forest in Sussex. Philosophical Transactions of the
Royal Society of London 115, 179-186.
Martin, P.S. and Klein, R.G. (1984). ‘Quaternary
extinctions. A prehistoric revolution’. (The University
of Arizona Press: Tucson, Arizona).
Mulvaney, J. an Kamminga, J. (1999). ‘Prehistory of
Australia’. (Allen and Unwin: Sydney).
Murray, P.F. (1978). Australian Megamammals:
Restoration of some Late Pleistocene fossil
marsupials and a monotreme. The Artefact 3, 77-99.
Murray, P.F. (1984a). “Australia’s prehistoric animals’.
(Methuen Australia: North Ryde).
Murray, P. (1984b). Extinctions downunder: a bestiary of
extinct Australian late Pleistocene monotremes and
marsupials. In ‘Quaternary extinctions. A prehistoric
revolution’ (Eds P.S. Martin and R.G. Klein) pp.
600-628. (The University of Arizona Press: Tucson,
Arizona).
Murray, P. (1986). Propalorchestes novaculacephalus gen.
et sp. nov., a new palorchestid (Diprotodontoidea:
Marsupialia) from the middle Miocene Camfield
Beds, Northern Territory, Australia. The Beagle.
Occasional Papers of the Northern Territory Museum
of Arts and Sciences 3, 195-211.
Murray, P. (1990). Primitive marsupial tapirs
(Propalorchestes novaculacephalus Murray and P.
ponticulus (Marsupialia: Palorchestidae) from the
middle Miocene Camfield Beds, Northern Territory,
Australia. The Beagle. Occasional Papers of the
Northern Territory Museum of Arts and Sciences 7,
39-51.
Murray, P.F. (1991). The Pleistocene megafauna of
Australia. In “Vertebrate Palaeontology in Australasia
(Eds P. Vickers-Rich, J.M. Monaghan, R.F. Baird and
T.H. Rich) pp. 1071-1163. (Pioneer Design Studio:
Melbourne).
Murray, P.F. and Chaloupka, G. (1984). The Dreamtime
animals: extinct megafauna in Arnhem Land rock art.
Archaeology in Oceania 19, 105-116.
Outram, D. (1984). ‘George Cuvier. Vocation, science and
authority in post-revolutionary France’. (Manchester
University Press: Manchester).
Owen, R. (1841[1842]). Report on British fossil reptiles:
Part Il. Report of the British Association for the
Advancement for Science (Plymouth Meeting). Pp.
60-204.
>
Proc. Linn. Soc. N.S.W., 130, 2009
Owen, R. [In Anon.] (1873). On the fossil mammals of
Australia. Family Macropodidae. Genera Macropus,
Leptosiagon, Procoptodon and Palorchestes. — Part
IX. Proceedings of the Royal Society of London 21,
386-387.
Owen. R. (1874). On the fossil mammals of Australia,
Part IX. Family Macropodidae: Genera Macropus,
Leptosiagon, Procoptodon and Palorchestes.
Philosophical Transactions of the Royal Society of
London 164, 783-803.
Owen, R. (1876). On the fossil mammals of Australia,
Part X. Family Macropodidae: mandibular
dentition and parts of the skeleton of Palorchestes;
additional evidence of Macropus titan, Sthenurus
and Procoptodon. Philosophical Transactions of the
Royal Society of London 166, 197-226.
Owen, R. (1877). “Researches on the fossil remains of the
extinct mammals of Australia; with a notice of the
extinct mammals of England’. (J. Erxleben: London).
2 volumes.
Owen, R. (1880a). Description of a portion of mandible
and teeth of a large extinct kangaroo (Palorchestes
crassus Ow.) from ancient fluviatile drift,
Queensland. Transactions of the Zoological Society
of London 11, 7-10.
Owen, R. (1880b). XXIII. Description of some remains of
the gigantic land-lizard (Megalania prisca, Owen),
from Australia. — Philosophical Transactions of the
Royal Society of London 171, 1037-1050.
Owen, R. (1894). “The Life of Richard Owen’. (John
Murray: London). 2 volumes.
Pinch, T. (1985). Towards an analysis of scientific
observations: the externality of evidential significance
of observational reports in physics. Social Studies of
Science 15, 3-37.
Piper, K.J. 2006. A new species of Palorchestidae
(Marsupialia) from the Pliocene and early Pleistocene
of Victoria. Alcheringa Special Issue 1, 281-294.
Quirk, S. and Archer, M. (1983). ‘Prehistoric animals of
Australia’. (Australian Museum: Sydney).
Ramsay, E.P. [In Anon.] (1885). Untitled. The Sydney
Morning Herald 14878, 10.
Raven, H.C. and Gregory, W.K. (1946). Adaptive
branching of the kangaroo family in relation to
habitat. American Museum Novitates 1309, 1-14.
Rich, P.V., van Tets, G.F. and Knight, F. (1985).
‘Kadimakara. Extinct vertebrates of Australia’.
(Pioneer Design Studio: Melbourne).
Rich, T. H. (1991). Appendix II: Literature References
to the Fossil Terrestrial Mammals of Australia.
In “Vertebrate palaeontology in Australasia’ (Eds
P. Vickers-Rich, JM. Monaghan, R.F. Baird and
T.H. Rich) pp. 1058-1069. (Pioneer Design Studio:
Melbourne).
Rich, T.H., Archer, M., Hand, S.J., Godthelp, H.,
Muirhead, J., Pledge, N.S., Flannery, T.F.,
Woodburne, M.O., Case, J.A., Tedford, R.H.,
Turmbull, W.D., Lundelius, E.L. Jr., Rich, L.S.V.,
Whitelaw, M. J., Kemp, A. and Rich, P.V. (1991).
Appendix I. Australian Mesozoic and Tertiary
terrestrial mammal localities. In “Vertebrate
35
RECONSTRUCTING PALORCHESTES
palaeontology in Australasia’ (Eds P. Vickers-Rich,
J.M. Monaghan, R.F. Baird and T.H. Rich) pp. 1005-
1058. (Pioneer Design Studio: Melbourne).
Roberts, R.G., Flannery, T.F., Ayliffe, L.K., Yoshida, H.,
Olley, J.M., Prideaux, G.J., Laslett, G.M., Baynes, A.,
Smith, M.A., Jones, R. and Smith, B.L. 2001. New
ages for the last Australian megafauna: continent-
wide extinction about 46 000 years ago. Science 292,
1888-1892.
Rudwick, M.J.S. (1992). ‘Scenes from deep time: early
pictorial representation of the prehistoric world’.
(The University of Chicago Press: Chicago).
Ruhen, O. (1976). “The day of the Diprotodon’. (Dai
Nippon: Hong Kong).
Rupke, N.A. (1994). ‘Richard Owen. Victorian naturalist’.
(Yale University Press: New Haven).
Scott, H.H. (1915). A monograph of Nototherium
tasmanicum (Genus - Owen” Sp. nov). Zasmanian
Department of Mines Geological Survey Records 4,
1-46.
Scott, H.H. (1916). A note of “Palorchestes” as
a Tasmanian Pleistocene genus. Papers and
Proceedings of the Royal Society of Tasmania 1924,
53-58.
Simpson, G.G. (1945). The principles of classification and
a classification of mammals. Bulletin of the American
Museum of Natural History 85, 1-350.
Stirling, E.C. and Zietz, A.H.C. (1900). Description of the
bones of the manus and pes of Diprotodon australis,
Owen. Memoirs of the Royal Society of South
Australia 1, 1-40.
Stirton, R.A. (1957). Tertiary marsupials from Victoria,
Australia. Memoirs of the National Museum of
Victoria 21, 121-134.
Stirton, R.A. (1967). The Diprotodontidae from the
Ngapakaldi Fauna, South Australia. Bureau of
Mineral Resources, Geology and Geophysics Bulletin
85, 1-44.
Tate, G.H.H. (1948). Studies on the anatomy and
phylogeny of the Macropodidae (Marsupialia).
Results of the Archbold expedition, No. 59. Bulletin
of the American Museum of Natural History 91, 237-
85k
Tedford, R.H. (1991). Vertebrate paleontology in
Australia: The American contribution. In ‘Vertebrate
palaeontology in Australasia’ (Eds P. Vickers-Rich,
J.M. Monaghan, R.F. Baird and T.H. Rich) pp. 46-83.
(Pioneer Design Studio: Melbourne).
Trezise, P. (1971). ‘Rock art of south-east Cape York’.
(Australian Institute of Aboriginal Studies: Canberra).
Van Reybrouck, D. (1998). Imaging and imagining
the Neanderthal: the role of technical drawings in
archaeology. Antiquity 72, 56-64.
Vickers-Rich, P.V. and Archbold, N.W. (1991). Squatters,
priests and professors: A brief history of vertebrate
palaeontology in Terra Australis. In ‘Vertebrate
palaeontology in Australasia’ (Eds P. Vickers-Rich,
J.M. Monaghan, R.F. Baird and T.H. Rich) pp. 1-44.
(Pioneer Design Studio: Melbourne).
36
Wells, R.T. (1975). Reconstructing the past. Excavations
in fossil caves. Australian Natural History 18, 208-
Ae
Wells, R.T. (1978). Fossil mammals in the reconstruction
of Quaternary environments with examples from
the Australian fauna. In ‘Biology and Quaternary
environments’ (Eds D. Walker and J.C. Guppy)
pp. 103-124. (Australian Academy of Sciences:
Canberra).
Whitley, G.P. (1966). Some early references to the extinct
marsupial Zygomaturus. Australian Zoologist 13,
228-231.
Williams, J. (1991). Dinosaurs - The first 150 years. A
brief history of dinosaur discovery. Geoscientist 1,
19-22.
Woodburne, M.O. (1967). The Alcoota fauna, central
Australia. An integrated palaeontological and
geological study. Bulletin of the Bureau of Mineral
Resources 87, 1-187.
Woods, J.T. (1958). The extinct marsupial genus
Palorchestes. Memoirs of the Queensland Museum
13, 177-193.
Wroe, S., Field, J., Fullagar, R. and Jermin, L.S. (2004).
Megafaunal extinction in the late Quaternary and the
global overkill hypothesis. A/cheringa 28, 291-331.
Yunupingu, G. (ed.) (1997). ‘Our land is our life. Land
rights — past, present and future’. (University of
Queensland Press: St Lucia).
Proc. Linn. Soc. N.S.W., 130, 2009
A Basal Actinopterygian Fish from the Middle Devonian
Bunga Beds of New South Wales, Australia.
BRIAN CHOO
School of Earth and Marine Sciences, The Australian National University, Canberra, ACT 0200, and
Museum Victoria, PO Box 666E, Melbourne, Victoria 3001 (mail correspondance to latter address)
bchoo@museum.vic.gov.au
Choo, B. (2009). 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. Linn. Soc. N.S.W., 130, 2009
P.J. MYERSCOUGH
I I I G cE Os Og Cade L
if 14 ¢ vl 69 IdeL
8C LE OV 8V L9 Ch Vell €dcl
€l 91 v6 CUCL
8 8 Il Il II I] II Il I] I] I] IO. Stl 691 COC 6th OCh Ltr CM EL
6 6 6 6 6 Cl él 81 81 81 ral 98 i oils Gs Wir Wwe WAC IME
L fl iL L fl fl (L Ol Ol Ol Ol €V 6V 09 v8 99 9LC LBC EMCL
€ 6 6 Cl cl 66 Stel WAN SOE SOG EG ESC IMCL
py ofisuo]qo g
Ol Ol Gl Cl Cl Gil tall €l vl Vall vi LI LI ial LI vv Iv 6S cae L
I I it G € € v Ol 9 cv IGEL
Ol Ol Cl Cl Cl Cl Gl cl Gl Gil Gl vv OV LV 8 VS igs VS €dcl
I I I It I I C C G C C Gl LI {| LI CC GG CS CaCL
VA VV cv LV LV LV SV 8V 8V OS IS 19 19 19 (69) 89 $9 teil CMEL
Gil vl 8I OC OC Ke Ic GG GC €C €C Of O£ O€ [€ Of CE CE [MEL
9 9 9 9 9 9 9 9 9 9 9 6 6 6 Ol LV 4 LV EMCL
V ¢ L 6 Ol cl Cl Cl tall €l el Ic IC GG CC LE 6 VC IMCL
pinwap “g
if I IL €CC cde L
I € 68 IGEL
€ (G0) cdc
I G 0cI CUCL
06 Icl 8Il Lcel OE COE 096 CMEL
Ne €9 06 O€l coc Otc LOD IME
ce 6£ OS C8 CLI 9SC SOS EMCL
8V 8S L8 LOI Sc O06C LOY IMCL
ppunqisoyf ‘q
I It I I € € € € € iS ¢ ¢ Cl vi cl SS Cae
¢ ¢ 8 8 8 8 Il Cl Cl el el VI VI VI 6l 6 6c CS IGE
Ol Ol Il el Li LI eV €dcL
C G C v Ol VI eV CUCL
I I I V CC VA 99 CMEL
€ (e V All Of Se OL [MEL
Ol Ol Ol LI CC 6£ (6S EMCL
€ ¢ [l GG €€ Gu VS IMCL
pyofiayn y
SS) OMS EES IES A SIS tell SCI IOI OS OL BOG SS Wy YS) OSS Sin iE MOG CS Tar out
"1661 YAR] UL SUIMOS JdUIS (SAvdA) BUNTY YSNOAY) Spor [eJUIUITAIdX9 UO SUTATAINS SSUI[ Pes JO JOAQUIN *¢ 9IqUL
53
Proc. 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.
Keith, D.A., Tozer, M.G., Regan, T.J. and Regan H.M.
(2007b). The persistence niche: what makes it and
what breaks it for two fire-prone species. Australian
Journal of Botany 55, 273-279.
Myerscough, P.J. and Carolin, R.C. (1986). The
vegetation of the Eurunderee sand mass, headlands
and previous islands in the Myall Lakes area, New
South Wales. Cunninghamia 1, 399-466.
Myerscough, P.J. and Clarke, P.J. (2007). Burnt to blazes:
landscape fires, resilience and habitat interaction in
frequently burnt coastal heath. Australian Journal of
Botany 55, 91-102.
Myerscough, P.J., Clarke, P.J. and Skelton, N.J. (1995).
Plant co-existence in coastal heaths: floristic patterns
and species attributes. Australian Journal of Ecology
20, 482-493.
Myerscough, P.J., Clarke, P.J. and Skelton, N.J. (1996).
Plant co-existence in coastal heaths: habitat
segregation in the post-fire environment. Azstralian
Journal of Ecology 21, 47-54.
Pate, J.S., Froend, R.H., Bowen, B.J., Hansen, A. and
Kuo, J. (1990). 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. EA ieH bik TU BREOM Hier ntlbe sy
ORR RAMURY SIRNA e ar aA bind ee.
t Shs yal!
sepulert palate hon a PANE
Aysabet, wcrrt cote aati Bo
write ala ipo. wot No ea
Nunbie dened seh iy
pret amare SW! a
{" re a diy attic ap
se! W Conihel Mu, AAO)
bint scala ab ies
| SS: CHIBI eT
‘be MWasirag to pmay eno T FEA ly, ay ity Ae
tah | SP Ra caved TRG Si
io Chee iid rind tub pene Sia ete
joe's ES? 7B ots
ihe AKU Pela bd aod? ta 19%
Baprnk Seth tat bak eve oom &
NiCr S Bite area PR He Set ey:
‘ou tent OP
ide yal. A naa dvrenl iene
Hobie Rormiiil Micubs bag iia si8O: :
tise ks
i vier renin Ray ‘Henig sth |
Rd ee ner ow ADL Kt rat siren ala oe t ;
head Andries yy, J Sirah, © PE sicentont
was” eastern weaiibenrsnats
pnttegy ae i De eke sos. i
Hiner Pr nd p (iRcey pce a
WOR oie ine than DM sor, pie Si
tila abuts Fade Sigh ahah au SHE
kT Sitio tft Se HN aE ANS
Pcotoginyl Socie nod aig welt re et Hn
eRe Sead bind! eB caer Bae PERSE GR
walt anaes T Hiseke Sy el aban a Oe BAN
ea oS Td Malina NOL
dette: (roee FL SRE baw U9 fade
Fal aedheddy seth third Wd! storie
Ve Sire base iin OS RAS "Gah
st pas ty COeaernelared), cay. fiQ! 22 uorc8 | "a
Ciel! ictal ia ee ie aa
arsoniy iter Hilf Tacs oS Gs Aaa Stat (
ia Hedehiel, “Ania, eH ageRe Bat
‘SAHA fsa 7 Leb Chk Me
CHEMO Police ae. Ea aahar sores “Ab
ON arated Seas rae oaitll Aaah
oi i nn ‘ghee 4 ‘it SoH} cH ‘es pee
pret ®) pee pere spylesa \o. Yotirwtol
it ri tila. 8 Swe Rob soi Seema
OF ahd ili here HAR | FeO PNR a
Vel BST Seb (Bay hh a wMS SSG
abtichrier a! 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).
The Carboniferous dendroid graptolites of Britain
and Ireland. Proceedings of the Yorkshire Geological
Society 49, 295-319.
Edgecombe, G.D. and Sherwin, L. (2001). Early Silurian
(Llandovery) trilobites from Cotton Hill, near Forbes,
New South Wales. Alcheringa 25, 87-105.
Fowler, G.H. (1892). The morphology of Rhabdopleura
allmani. In R. Leukarts (ed.) Festschrift cum Toten
Gebiirstag, W.S. Engelmann, Leipzig.
Frech, F. (1897). Lethaea geognostica; Theil |, Lethaea
palaeozoica, | Band, Graptolithiden (Leif. 1-2 by F.
Roemer; Lief. 3 by F. Frech), 544-684. Stuttgart. E.
Schweitzerbart’sche.
Hall, J. (1843). Geology of New York, IV, Survey of the
Fourth Geological District. Natural History of New
York. Geology of New York, 156 pp.
Hall, J. (1851). New genera of fossil corals. American
Journal of Science 11, 398-401.
Hall, J. (1858). Descriptions of Canadian Graptolites.
Geological Survey of Canada, Report for 1857, \\\-
145.
Hall, J. (1862). New species of fossils from the
investigations of the Survey. Wisconsin Geological
Survey, Report for 1861, 1-18.
Hall, J. (1865). Graptolites of the Quebec Group. Canadian
Organic Remains, Geological Survey of Canada 2(i-iv),
151 pp.
Harmer, S.F. (1905). The Pterobranchia of the
Siboga Expedition, with an account of their
species. Uitkomsten op zoologisch, botanisch,
oceanographisch en geologisch gebied verzameld in
Nederlandsch Oost-Indié 1899-1900 aan boord H.M.
Siboga 12, Monographie 26, 132 pp. Leiden.
Lankester, E.R. (1877). Notes on the embryology and
classification of the Animal Kingdom: comprising
a revision of speculations relative to the origin and
significance of germ layers. Quarterly Journal of
Microscopical Science 17, 399-454.
Lapworth, C. (1881). On the Rhabdophora (Hopk.) or
dendroid graptolites collected by Prof. Keeping in the
Llandovery rocks of Mid Wales. Quarterly Journal of
the Geological Society of London 37, 171-177.
Loydell, D.K., Storch, P. and Melchin, M.J. (1993).
Taxonomy, evolution and biostratigraphic importance
of the Llandovery graptolite Spirograptus.
Palaeontology 36, 909-926.
Miller, S.A. (1889). North American geology and
paleontology. Western Methodist Book Concern,
Cincinnati, Ohio, 664 pp.
Nicholson, H.A. (1872). Monograph of British graptolites,
i-x, 133 pp. Blackwood and Sons, Edinburgh,
London.
76
Packham, G.H. (1967). The occurrence of shelly
Ordovician strata near Forbes, New South Wales.
Australian Journal of Science 30, 106-7.
Pocta, P. (1894). Systéme Silurien du Centre de la Bohéme,
8, pt. 1, Bryozoaires, Hydrozoaires et parte des
Anthozaires. Prague, 230 pp.
Prout, A.H. (1851). Description of a new graptolite found
in the Lower Silurian rocks near the Falls of St. Croix
River. American Journal of Science 11, 187-191.
Rickards, R.B., Chapman, A.J., Wright, A.J. and Packham,
G.H. (2003). Dendroid and tuboid Graptolites from
the Llandovery (Silurian) of the Four Mile Creek
Area, New South Wales. Records of the Australian
Museum 55, 305-330.
Rickards, R.B. and Durman, P.N. (2006). Evolution of
the earliest graptolites and other hemichordates. In
Studies in Palaeozoic Palaeontology, M.G. Bassett
(ed.). National Museum of Wales, Geological Series
25, 5-92.
Rickards, R.B., Hamedi, M.A. and Wright, A.J. (2001).
New assemblages of graptolites, rhabdopleuran
hemichordates and chitinous hydroids from the
Arenig (Ordovician) of the Banestan area, Iran.
Alcheringa 25, 169-190.
Rickards, R.B., Packham, G.H., Wright, A.J. and
Williamson, P.L. (1995). Wenlock and Ludlow
graptolite faunas and biostratigraphy of the Quarry
Creek district, New South Wales. Association of
Australasian Palaeontologists 17, 68 pp.
Rickards, R.B. and Wright, A. (1997). Graptolites of the
Barnby Hills Shale (Silurian, Ludlow), New South
Wales, Australia. Proceedings of the Yorkshire
Geological Society 51, 209-227.
Rickards, R.B., Wright, A.J. Morgan, E.J. and Farrell, J.R.
(2005). Silurian graptolites from the Barnby Hills
Shale and Hanover Formation, New South Wales,
Australia. Proceedings of the Linnean Society of New
South Wales 126, 153-169.
Sherwin, L. (1970). Preliminary results on studies of
graptolites from the Forbes district, New South
Wales. Records of the Geological Survey of New
South Wales 12, 75-6.
Sherwin, L. (1973). Stratigraphy of the Forbes - Bogan
Gate district. Records of the Geological Survey of
New South Wales 15, 47-101.
Sherwin, L. (1974). Llandovery graptolites from the
Forbes district, New South Wales. Special Papers in
Palaeontology 13, 149-175.
Sherwin, L. (1976). The Secrets section through the
Cotton Beds north of Parkes. Geological Survey of
New South Wales, Quarterly Notes 24, 6-10.
Spencer, J.W. (1878). Graptolites of the Niagara
Formation. Canadian Naturalist 8, 457-463.
Proc. Linn. Soc. 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 <p Zone
hi =e fe
L cy 3 ai
20 i aS r
FT EI ee
i i Me ee
Ley patrol tt We real
Ray ty eg 5
40 | oR I i=
at . - L A
| |
60 C |
fur Pt i;
int
/
|
L
|
sun
Figure 4 continued
European part at the top of the profile. There would
have been a greater biomass after 2 cal ka, hence more
fuel to burn, particularly on the swamp itself.
Notts Swamp
Notts Swamp, at 33° 48’ 35.44” S, 150° 24’ 27.66”
E and about 682 m altitude is located in a shallow
hanging valley. Below the swamp, Reedy Creek
flows over a small cliff and follows a steep, narrow
valley into the Kedumba Valley. The Wentworth Falls
Claystone Member outcrops near the base of the
swamp.
Proc. Linn. Soc. N.S.W., 130, 2009
The lower third of the swamp is used for a
market garden, but there is no sign of disturbance or
weed invasion on the upper part of the swamp used
for this study. 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.
Stratigraphy: The core recovered 130 cm of
sediment. There was dark brown and greyish brown
peat with roots down to 50 cm, then black or very
dark greyish brown clay at 60-100 cm, with dark
grey or light grey sandy silt at 110-130 cm. Pollen
was recovered throughout the sequence, sometimes
in very high concentrations. Radiocarbon ages are
given in Table 4.
The swamp vegetation and surface pollen: Species
of Kunzea, Gahnia and Leptocarpus tenax were
dominant on the swamp. Species of Gleichenia,
Selaginella, Leptospermum, Cyperaaceae, Juncaceae
and a number of sclerophyllous shrubs were also
present (Chalson and Martin, this volume). In the
surface samples, Myrtaceae, Casuarinaceae and
Restionaceae were well represented. There was
appreciable Pinus pollen also. (Chalson and Martin,
this volume).
The pollen record: The pollen spectra from the
sediments are presented in Fig. 6A, 6B and is zoned
thus:
Zone D, 110-130 cm, c. ?7,300-4,500 cal yr BP
(for estimated ages, see Fig. 7). Myrtaceae pollen
content was low and Casuarinaceae moderate. The
Selaginella spore content was appreciable at the
base, decreasing through the zone. The Restionaceae
and Poaceae content was moderate and the lowest
for the profile.
Zone C, 100-80 cm, c. 4,500-2,400 cal yr BP. The
Myrtaceae, Restionaceae and Poaceae representation
increased but the Selaginella content was much
reduced when compared with the zone below, and this
change coincided with a change in sediments to clay.
Gleichenia and other fern spores increased somewhat
when compared with the zone below.
Zone B, 70-30 cm, c. 2,400 cal yr BP. to ?modern.
There were more identifications of the mytaceous
pollen, an increase in Restionaceae and few Gleichenia
and other fern spores when compared with the zone
below. The Se/aginalla and Cyperaceae content was
minimal.
87
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
50
100
Depth (cm)
150
200
250 - -
| : a Age (k yr) ‘
Radiocarbon date
Figure 5. Warrimoo Oval Swamp summary diagram.
Zone A, 20-0 cm, modern. Pinus was found throughout
the zone, indicating post European settlement. The
Myrtaceae and Casuarinaceae pollen content was
maintained. Restionaceae decreased towards the
top and the Cyperaceae content, although low, is
the greatest for the profile, when compared with the
zones below
The charcoal content was very low at the base
of the profile when Se/aginella was prominent on the
swamp, then increased after the decline in Se/agiella
and was consistently high in the European zone.
History of the Vegetation’ About 7-4.5 cal ka,
Selaginella was common on the swamp and the
surrounding vegetation was an open woodland, with
Casuarinaceae prominent. Fire was not common
then. After about 4.5 cal ka, Se/aginella was replaced
Table 4. Radiocarbon ages for Notts Swamp
@ Calibrated date
by Restionaceae, the tree
cover increased somewhat
and fire became more
common. The vegetation
remained relatively stable
until modern times when
there was a slight decrease
in Restionaceae and an
increase in Cyperaceae.
Charcoal abundance was
higher when the tree cover
was greater.
European Zone
Ingar Swamp
Ingar Swamp, at 33° 46’
165: Sxl S02 Die 2229 22a:
and 584m altitude, is broad
with many channels and
hummocks of Cyperaceae
forming ridges. The Banks
Wall Sandstone Formation
underlies the swamp and
4 there are outcrops of the
Wentworth Falls Claystone
Member near the lower
margin of the swamp. The
swamp occupies the floor of
a Shallow hanging valley on
the plateau surface. Below the swamp, Ingar Creek
forms a waterfall where the valley gradient steepens.
Stratigraphy: The core recovered 155 cm of sediment.
Peat with roots was found at 0-20 cm, then humic clay
with roots at 25-110 cm, sandy humic clay at 120-130
cm, then sandy clay at 135-145cm, and silty clay at
150-155 cm. The radiocarbon ages are given in Table
3s
The swamp vegetation and surface pollen: Species
of Leptospermum, Cyperaceae and Restionaceae
were dominant on the swamp. Gleichenia and
some sclerophyllous shrubs were also present
(Chalson and Martin, this volume). The surface
samples (Chalson and Martin, this volume)
showed that most of the Myrtaceae pollen was
: Radiocarbon years (yr Calibrated age (cal.
Depth (cm) Material dated Laboratory no. BP) yt BP)
25-35 Peat with roots SUA 2653 1.013+0.008x modern Modern (<33)
75-85 Clay SUA 2654 2,400+£70 2,578+130
120-130 Sandy silt SUA 2655 5,630+70 6,478+80
88 Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
‘xipuaddy aas ‘u0e}a80A ay) ul oA} uatfod a4) Jo 99.1n0s a[quqoad 104 ‘v.Qd0ds udT[od dues S}JON “G9 “V9 SSaANIY
J
io or Ae 1x8 OL X OL %0Z
sjoo Kel cael HIS [74] pues |:\'- weg |e So/SulEJB
= ral a — ig Sas NPB ae feraneeed jz
Sie bevad a =f siE9S
er |
a OstaZp'9 ‘a oe ; :
oF ozt
oe “ ‘a eee ieee ie ans See RE TER is
a OOL
is 2
Paine og
— OfLFasc'z
yee
cee 09
5) 5
Se v
AY Ov
LMT
. Wwspoy
Fo fie Aw
: 0¢
ray
¥Y| auez
Ba 2d ce a T 7 7 ones 0
sg b P L r
& jeoo1eyo = ~ £
a aidossoJoeyy ao ah =
ts) eo
— > ¢
v
a
f V9
89
Proc. Linn. Soc. N.S.W., 130, 2009
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
ponulyjuod 9 sins
Depth (cm)
hh —
wn So oO
° ° fon) ro)
Oo
=
wie
re
E
3 ra
ny oO
z =
g a
S Es
=
<4}
BA
=
tS}
=)
s =~
i“)
=
=)
= o—
) A
= e)
“< o 8
a gz 8
Fy a 3 o
= 2. 3
iS) e. =!
5 os
a
90 Proc. Linn. Soc. N.S.W., 130, 2009
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
&
& é
ae SB hig@ak
ef e é
& é av & @ & £ f e oO & e —
~— é o B) FS 1
ceded? ¢G |b & LF Gb us §f
See ds £ Ss F Q & sé & fou 4 ris se
fy : = —— - me pil a
207 x
2 - : ; hoe
Modern
a
40
60
80 B
100
420 4 ue —
7,428490 —'
ike
be 7,188+90 “_-
ets
= waee
cont)
Ss Peat
- Aw
Figures 8A 8B. Ingar swamp pollen spectra. For probable source of the pollen type in the vegetation,
see Appendix.
Scale
ram font
20% 10X10" grains/cc
Proc. Linn. Soc. N.S.W., 130, 2009
ES Fe E |
~'s\"| Sand Silt | Clay M Roots
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
‘x1pusddy aas ‘u01e}030A ayy Ut 9dA} UDTJ[Od 94} JO 9d4N0s a[quqoad 104 “v.aQd0ds Uatjod dulEMG equiooyey “G71 “VZI SdINSI]
s}ooy gy Kelp ae WS Z ] pues }e9
“, OLXShy ° sions 7 ve ceeence cert nau manila ay a eens on | | |
od1#8s6" rAUAE | Se MKB esseenezey 01 x8h: Sail Tee os - 2 | ree
Sw val SOLK Lg -- ee PO hae reek oe see $e dere : | | ‘ Or
ts vente QEMQZ cree meee ece seach vette esos ne eae camel geteeee Ty te remem el). ga Sl RE et eet et ce
; al ae ytie ee eceeme peer as wien ‘ gay pOb XOL 2
SE OLKGE SF yOLXOTT - wes a - vm mare x y Fat ae
1) a] aes nrenees <3 OLX9CP + “eae x seer Benes apse DS hea | * 02
aed AE ODMR to Se ey orue wea eeeecestrgd ahve vate neon cmos mena ie trices sean Bae wrsainhd auwism ace j;—_" minsins
ai at [| 0 gaa OP RA Sma Cetae bee ree oases ajeos a
—! if ip ipcatraciaaed i =, 0LXG¢ Be S ghy po onteritan as: raat oH | 0
OZ1LF866 cl
ieee
o0L#89z'9
weap d pODXP PL sonm sae oe macnn nd moe a eee pee ie ae
2 ctanseh we pabeeet ii 5k od bon rarer com | POD Kit anes tease A= Sees peeeer wee © 7 ra WA So/SuIEUB ,OL x OL
= eg rie oe a es ees
Ey Oz
—| 5 ,OLXOS! we es se Zing neue eee mee aremeg » -
ammo rid OLX PEP mci eesse aes] PL OLXZO) “on “Ae ee ~
eeeren Lx Bea sek ees ord |
panel <,0.x£0P eR arp Re Py 01xoel ceeetatiser asterd <3, OLXG'9 = { OL
ac OLXQT) emcee ate) A nee ee oem EY
cease = 0Lxoge=~ “= maw ee etn] SO LNOL | asses mene om een OL XSL. rena =, %0C
Ss —w ; ajeas
Pol hoy eee eee |e aPertcre mec aes 3.0
aac, rye Cae
pOLXCOZ -—- « pe cases j§,0LXBZL Ap 0s wereae rou hans ad weiss eee nee see | q onan
09
ov
_—— ~n ee rer ce i myn en
— = rie > Nir moe pores cet tk oz
i Nee OLX £97 “nese nese == oe
an =|4,OLXP = 24 ,0XEe2 SS —_
saa Se ane pce ae | AYO J
oa,
fBooreyo
didossoJoEyy
97
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
Ys oth | . 8
Sera er a ee
o oO Oo oS
i= o ve oO
Sot 3 ee oT ok
es 2 a rs a >
if e _ ae ra)
8 j 7 =
eS | If “ip o
BS rh bee, eg, 8 < a8] ) a
| | | | | : By, : a
Sa, r - — :
00) | c=) Ou.
= Uig Soy Sy oH
Yay ss Mo, =
10 : Me
oe SS
of jen E Mf :
“% ; [ re Arey hy y
o > “a5
= * oe b secret oereore as
: ih i [= ES Vee, pes
Pop, ‘ h ! i t E i Sy, <i oe ; Mes
Gye teeta dt EB Ce,
sy |i lit UE ae Ea Hi a Mes ue!
B: fay
% i ‘/a) es 0 iS ar ae ar eS | IB
roi! &
i E Fe,
fae YL
ey : 7 e a8 Ze
3 alta: : erg Vn,
ay Qe, “YW
: 1 Ao.
Tf “oy B= (8 =e
‘ feel) ete @) at
pk 4 Lub beun 9 aon
fi et 7 86. #77)
d Fi » ot i H i = Pay Y
Wi, Ei i g & 40,
oy Wty, 49yy
; . i e TS Corn Jo eee (eh +
ih Sos
ro i iN 4 a) . at. oo
: ' xv 2
&, [=]
— faa) ®2p, a
s Tt Kay, x
=) ™
os oly i) o o o =)
@ ? N + © a
8
w
i ®
Proc. Linn. Soc. N.S.W., 130, 2009
Figures 14A, 14B. Newnes Swamp pollen spectra. For probable source of the pollen type in the vegeta-
100
tion, see Appendix.
J.M. CHALSON AND H.A. MARTIN
Depth (cm)
on
o
Zone D @ \--2:
100+——— = . beds Bis
0 2 4 8 10 12
Age (k yr)
I Radiocarbon date
@ Calibrated date
Figure 15. Newnes Swamp summary diagram.
History of the vegetation: The vegetation was open
Eucalyptus woodland at 11 cal ka, but by about 7.5 cal
ka, Eucalyptus species had disappeared, Casuarinaceae
and the sclerophyllous shrubs increased, suggesting a
heathland. After 2 ka, Melaleuca became prominent,
possibly around or on the swamp. Burning was
moderate to low in the early Holocene, very low in the
mid Holocene when the vegetation was a shrubland
or heath and after 2 ka, it was moderate to high, when
Melaleuca had colonised the swamp.
DISCUSSION
Stratigraphy
All of the swamps chosen for this study are found
associated with small streams in valleys of the rugged
terrain of the Blue Mountains. While such sites
may not be the first preference for palynology, they
allow study in an area where the more favoured sites
are rare. These small valley swamps rely on some
barrier, often a clay substrate, to impede drainage and
maintain the swampy conditions. The root mats of
the vegetation stabilise the sediments and slow down
the water flow, but if the vegetation is disturbed, then
the sediments are prone to erosion. The swamps dry
out occasionally but not seasonally. The vegetation
can withstand mild or short droughts, but prolonged,
severe droughts such as has been experienced in
recent years destabilise the communities as some
species die and others replace them. The swamps then
become very vulnerable to fire, human trampling or
even the next major rainfall event. Elimination of the
vegetation cover over even a small area of the swamp
leaves it vulnerable to subsequent erosion.
If the vegetation is destroyed and there is erosion,
channelised water and higher energy flows deposit
coarser grained sediment, such as sand. Eventually
Proc. Linn. Soc. N.S.W., 130, 2009
the vegetation re-establishes and stream flow slows
down and finer particles, such as silt and clay are
deposited.
It has been assumed that the sediments were
deposited at a uniform rate: however, the resolution
of dating does not allow this to be tested. Uniform
rates of sedimentation are probably not the case at
finer scales of resolution.
The peat layer at the top of the swamp is usually
only 20 cm or less in thickness. While roots of the
present vegetation may penetrate to a considerable
depth, a discrete layer with roots at depth in some
profiles suggests former peat or vegetation layers that
have been buried, and the decay of most of the organic
matter as the sediments accumulated. Also, there may
be an appreciable humic content of the sediments at
depths in the profile, a further indication of decayed
vegetation.
Using the above description of the dynamics
of the swamps, the sediments are interpreted as
follows:
Burralow_ Swamp: There is only some 1.2 cal ka
represented here, with sand at depth, then grading to
clay and peat at the top. The rate of accumulation of the
sand was rapid, with the clay and peat accumulating
much slower (from Fig. 3). It is likely that the whole
of this profile post-dates an erosive event.
The two basal two dates are puzzling, given that
they do not conform to the uniform sedimentation
rate discussed above. They are within the sand
layer, which was carbon poor, and it is possible
that groundwater carrying humic acids could have
contaminated the sediments with younger organic
matter, overwhelming the small quantities of older
carbon.
101
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
Warrimoo Oval Swamp: A basal sand layer dating to
about 4.7 cal ka grades into sandy silt, then another
sand layer at about 1.2-1.5 cal ka. The sediments then
became increasingly peaty towards the top. Deposition
of the basal sand layer probably followed an erosive
event, and the sand layer at 50-90 cm probably
represents another erosive event. This latter layer
may correlate with the basal sand layer in Burralow
Creek (from Figs 3 and 5), but this hypothesis requires
additional dating control to test it.
Notts Swamp: About 7 cal ka are recorded here (Fig.
7). The basal sandy silt layer is overlain by clay, with
peat with roots above it. The profile appears to reflect
a low energy depositional environment throughout.
The stratigraphy suggests that the lower and upper
layers may have accumulated at a somewhat faster
rate than the clay in the middle.
Ingar Swamp: This profile also represents about 7 cal
ka (Fig. 9). Sandy clay formed the basal sediments,
with clay with roots above it, then peat with roots
forming the top most layer.
Kings Tableland. Over 10 cal ka, the majority of
the Holocene is represented here (Fig. 11). There is
a basal sand layer, then a complex stratigraphy of
clay, sand, silty peat, sand and sandy peat above it.
This suggests that conditions of deposition would
have fluctuated, and in which case is unlikely that the
sequence is continuous.
Katoomba Swamp. Over 6 cal ka are represented in
core | and 10-12 cal ka in core 2 (Fig. 13). There are
no large sand layers similar to those seen in sediments
at some of the other sites, but a complex stratigraphy
of finer sediments, often with a sandy component.
Newnes Swamp. About 12 cal ka is recorded here
(Fig. 15). The sediments are sand then sandy or
silty clay in a complex stratigraphy at the base of
the profile. Above this, there is a prominent sand
layer, then sandy clay and peat with roots at the top.
Superficially, it appears that the sand layer in the
middle of the profile accumulated very slowly (Fig.
15), but another interpretation is possible. The date
at the top of this sand layer is about 1.3 ka, which
approximates the date of the top of the sand layers
seen in Warrimoo Oval Swamp and Burralow Creek
Swamp. If the sand layer does represent the aftermath
of an erosive event, then a section of the sediment
profile is likely to have been lost. The roots in the
sandy clay at the base of the sand layer may indicate
the base of a peat or vegetation layer that was buried
102
by the accumulating sand.
Each swamp thus has its own history of
sedimentation. Sandy layers in three of the swamps
suggest erosion after disruption of the vegetation,
sand deposition, then stabilisation sometime about
1.2-1.6 ka, with subsequent re-establishment of the
vegetation and deposition of fine-grained sediments.
If fire was the cause of this erosion, then we
could expect evidence of it in the charcoal record, but
there is no evidence of increased charcoal at this time.
Absence of charcoal cannot be taken as evidence of no
fire, as erosion may well have removed the charcoal,
along with some of the sediments. Fire is not the only
likely cause: as discussed, prolonged drought could
also destabilise these systems. Minor tectonics along
fault lines in the Blue Mountains (Bembrick et al,
1980) would also accelerate erosion.
The three swamps which have this sand body
are Newnes, Burralow Creek and Warrimoo Oval.
Newnes and Burralow Creek are the two most
northerly swamps and Burralow Creek and Warrimoo
Oval are the two most easterly swamps. Whatever
the cause of this disturbance, it seems to have come
from or been concentrated in the north east (see
Fig. 1). That Burralow Creek Swamp has only | ka
of sediment suggests that it may have suffered the
greatest disturbance and erosion.
History of the vegetation
The swamp vegetation. The survey of the vegetation
(Chalson and Martin, this volume), shows. that
species of Restionaceae, Cyperaceae, Gleichenia,
Selaginella, Baeckea, Kunzea and Leptospermum
dominate the vegetation cover of these swamps.
Many of the common sclerophyllous shrubs have
been recorded on the swamps, though not dominant,
as well as in the dryland vegetation (Chalson and
Martin, this volume). Poaceae has both dryland and
swamp species (Sainty and Jacobs, 1981).
In the pollen diagrams, Gleichenia and
Selaginella are found predominantly where the
sediments are sandy and Restionaceae is dominant on
the clayey sediments. There is very little Cyperaceae
here, unlike other sites, e.g Lake Baraba (Black et al.,
2007), Dry Lake, (Rose and Martin, 2007), Mountain
Lagoon (Robbie and Martin, 2007) and Penrith
Lakes (Chalson and Martin, 2008) which have more
Cyperaceae than Restionaceae. The swamps of this
study, however, are more ephemeral and unlike the
others with more Cyperaceae, which are lakes or
lagoons where the water would be more permanent.
Indeed, there are many species of Cyperaceae that are
aquatic (Sainty and Jacobs, 1981) whereas species of
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Restionaceae are found more in damp and swampy
places. Thus Cyperaceae flourishes in the more
permanently wet swamps and Restionaceae is more
abundant in these swamps subjected to irregular
drying. Species of both families may be found in the
dryland vegetation but the pollen record is heavily
dominated by the wetland species.
Baeckea, Kuzea and Leptospermum species are
present in the pollen diagrams of the swamps, but
mainly towards the top and especially in the European
zone. There is very little pollen of these taxa at depths
in the profiles. Some Melaleuca pollen is present and
it shows much the same trends. Although the trend
to more of these shrubs started before European
settlement, it appears that these woody shrubs,
which are often dominant on the swamps today,
have probably been further encouraged by European
activity, probably by the altered fire regime (Kohen,
1995).
The swamp vegetation thus reflects the sediment
substrate and hydrological conditions, with some
changes due to European activity.
The dryland vegetation. The sites are examined in
a time sequence to determine if there has been any
synchronous changes in the vegetation across the
Blue Mountains.
Three sites record the early Holocene of 10 ka
to 6 ka: Kings Tableland, Katoomba and Newnes
Swamps. About 10 cal ka, Eucalyptus species were
present at all three sites, but there was very little at
Kings Tableland. Casuarinaceae, the other group
which could be either trees or shrubs was present
also. Thus all three sites appear to have been wooded
in the early Holocene, with Kings Tableland probably
more open than the other sites. By 6-4 cal ka, the mid
Holocene, there were virtually no Eucalyptus in any of
the sites. The vegetation had become more open and
probably more of a sclerophyllous shrubland or heath.
Eucalyptus returned to the Katoomba site about 3 cal
ka, but very little is recorded in Newnes and Kings
Tableland up to the present. The Katoomba swamp
is located in a narrower and steeper valley than the
other two sites, and this shelter may have produced
better moisture retention and hence tree regeneration.
Melaleuca became established at Newnes about 1.3
cal ka.
Two sites date from about 6 cal ka, the mid-
Holocene: Notts and Ingar Swamps. Species of
Eucalyptus and Angophora were present at both
sites, hence they were probably wooded at the time
that Kings Tableland, Katoomba and Newnes were
dominated by shrubs. Warrimoo Oval dates from
about 4 cal ka, and the relatively low frequencies of
Proc. Linn. Soc. N.S.W., 130, 2009
Eucalyptus and Angophora indicate it was an open
woodland at that time.
In the period 4-2 cal ka, there was little change
from the previous period at Notts and Ingar Swamps.
At Kings Tableland, Casuarinaceae increased but there
was still no Eucalyptus. At Katoomba, Eucalyptus and
Angophora species reappeared, as this site probably
gained an overstorey of trees again. At Newnes, the
4-2 cal ka period was similar to that before, with
very little Eucalyptus. Burralow Swamp dated from
1 cal ka was initially very open, with the tree cover
increasing about 0.8 cal ka. Except for an increase in
Melaleuca or Leptospermum species in some of the
swamps, there was relatively few changes after 2 cal
ka until the European period,
In the European zone, there was minimal or no
decline in the Eucalyptus and Angophora content.
Casuarinaceae content declined noticeably at all
the sites. At Burralow, Warrimoo Oval and Kings
Tableland, the woody shrubs Callistemon, Baeckia,
Leptospemum and Melaleuca increased. The Poaceae
content remains unchanged in all swamps except for
Katoomba, where there is a dramatic increase.
There is thus relatively little change in the
palynology after European settlement in all of the
sites, except at Katoomba. This perhaps reflects the
relatively minor European changes to the sites, with
the exception of Katoomba where the swamp itself
has a history of use for various urban activities.
Agricultural development has been minimal, reflecting
the poor soils. The general lack of decline in tree
species is unexpected, but European development has
largely been confined to the ridgetops and extensive
natural vegetation is a feature of the Blue Mountains.
The wood of Casuarinaceae was prized by Europeans
as the firewood of choice for bakeries and the timber
had many uses (Entwisle 2005), hence it may have
been sought out more than the Eucalyptus species.
Each site has its own distinctive history, as
are the dominant Eucalyptus species at each site
(Chalson and Martin, this volume). There is limited
synchronicity of change between the swamps. The
three swamps at the highest altitude are the oldest,
dating to the beginning of the Holocene. They were
wooded in the early Holocene, but became very open
or almost treeless by the mid-Holocene. The sites at
the lower altitudes, however, were wooded during the
mid-Holocene. By the late Holocene, all of the sites
had become wooded, although the tree layer may
have been very open in some of them. Clearly, the
interplay of many environmental factors, not the least
of which is altitude, have influenced the vegetation at
each site.
Other sites in the Blue Mountains also present
103
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
unique histories when compared with those of this
study. At Mountain Lagoon (Robbie and Martin,
2007), the proportion of Casuarinaceae pollen is
substantial at the beginning of the Holocene, then
declines throughout the Holocene. A/locasuarina
torulosa 1s more common at Mountain Lagoon than
at any of the sites of this study. Pollen of swamp
plants increase through the Holocene at Mountain
Lagoon as the site developed from a lake in the early
Holocene to a peat swamp in the mid-late Holocene.
The Myrtaceae species identified are mainly different
to the species of this study and the proportion of pollen
remains much the same throughout the Holocene and
only declines after European settlement. The species
identified at Mountain Lagoon are often prized for
timber (Robbie and Martin, 2007). The physical
environment of Mountain Lagoon is totally different
to that of the Blue Mountain sites: it is a small basin
on Wainamatta Shale, in a particularly sheltered
location.
Kings Waterhole, part of the Mellong Swamps
in the Wollemi National Park, at 280 m altitude, has
a 6 ka history (Black and Mooney, 2007). Myrtaceae
(excluding Melaleuca spp.) and Casuarinaceae are
prominent until about 4-3 ka, when Casuarinaceae
begins to deline. At the same time, Restionaceae
increases. After 3 ka, there is minimal Casuarinaceae
and Restionaceae declines, but Melaleuca and Poaceae
increase. After 1 ka, Myrtaceae decreases somwhat
and Poaceae is prominent (Black and Mooney, 2007).
This decline of Casuarinaceae after 3 ka is not seen in
any of the sites of this study.
At Gooches Crater Swamp on the Newnes Plateau,
between 900 and 1,200 m altitude (Black and Mooney,
2006), there is a 14 ka history of the vegetation.
There is a moderate level of variability in the pollen
assemblages, and the swamp vegetation varied from
a wet heath with semi-permanent to permanent water
to a fern swamp. The Myrtaceae and Casuarinaceae
content is appreciable and continuously variable.
The Asteraceae content is considerable (Black and
Mooney, 2006), unlike the sites of this study, although
the Newnes site has the greatest Asteraceae content of
all the sites of this study.
Penrith Lakes on the Cumberland Plain just
east of the Lapstone Monocline has 6 ka of Holocene
history. The tree cover was very open in the mid
Holocene, becoming somewhat more wooded in the
late Holocene (Chalson and Martin, 2008), mirroring
the findings of this study.
The rugged terrain of the Blue Mountains would
have provided some isolation to each site so that
each has its own sedimentary and vegetation history.
Any climatic change or other regional event should
104
imprint in these deposits, especially in the more
environmentally sensitive sites.
Climatic change
The decline of the trees from the early Holocene
to the mid Holocene in the sites at the higher altitudes,
viz. Newnes, Katoomba and Kings Tableland, suggests
that the climate had become drier. A detailed analysis
of the climatic requirements of the Ezcalyptus species
also suggests a wetter early Holocene (Chalson,
199i):
Climatic trends in the mid Holocene are
uncertain, for while the sites at higher altitudes
were not wooded, sites at lower altitudes, i.e. Notts
and Ingar Swamps, were wooded at this time. Trees
returned to Katoomba about 3k yr, suggesting that the
climate had become wetter. As discussed previously,
the Katoomba catchment is narrower and steeper-
sided, hence the most sheltered of the three higher
altitude sites. Newnes and Kings Tableland, however,
remained open with few trees, suggesting that if
there was an improvement in the rainfall, it had not
returned to the early Holocene levels. These uncertain
trends continued into the late Holocene. About 2 kyr,
there was an increase in the wooded vegetation, with
more Eucalyptus at Warrimoo, more Casuainaceae
at Kings Tableland and more Melaleuca at Newnes.
The other sites, however, remained much the same.
There probably was an increase in rainfall, but it was
slight. The detailed analysis by Chalson (1991) came
to similar conclusions: climatic changes in the mid
and late Holocene are equivocal.
The climatic changes deduced from the this
study are in general agreement with other sites in the
Blue Mountains. The early Holocene is regarded as
a climatic optimum when it was warmer and wetter
(Allen and Lindesay, 1998). Evidence for the mid
and late Holocene is variable, some indicating wetter,
some drier conditions. Evidence suggests that the
El Nifio-Southern Oscillation (ENSO) phenomenon
came into operation about 5 ka, with increasing
seasonality. Thus from the mid Holocene, the climate
became more like that of today, with more variability
(Allen and Lindesay, 1998; Moy et al., 2002; Donders
et al., 2007).
Fire history
Charcoal has been found in all of the sites and
throughout all of the profiles. In the early Holocene,
the charcoal content was low to moderate in Kings
Tableland, Katoomba and Newnes. By mid Holocene,
the quantity of the charcoal had declined in Katoomba
and Newnes: there had been a change in the vegetation
from more wooded in the early Holocene to less
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
wooded in the mid Holocene, hence there may have
been less fuel to burn.
In the mid Holocene at Notts, Ingar and Warrimoo
Swamps, there is very little charcoal. All of the sites
would have been wooded to some degree but not as
much as in the late Holocene. The charcoal content
increases at each site as the tree cover increased.
In the late Holocene, charcoal content is variable,
but mainly greater than in the mid Holocene. In the
European zone, charcoal content is consistently high
and the highest for the profile, with the exception of
Katoomba. It may be that because of urban use of the
swamp at Katoomba, fire was excluded. Within these
trends, there may be the occasional single high value,
but they do not form any pattern.
The interpretation of a charcoal record is
problematical because so many factors are involved,
e.g. fire frequency, intensity and transport of charcoal.
The results of this study suggest that the greater the
biomass, the more fuel there is to burn hence the more
charcoal in the sediments.
The higher charcoal content of the European
period suggests that fire regimes were changed with
settlement. If Aboriginal people regularly burnt off
the undergrowth and suppressed the shrubs, then
the fuel load would be kept down. With European
settlement and the cessation of traditional fire
practices, it is possible that the woody shrubs
became more common (Kohen, 1995; Ward et al.,
2001). Under these conditions, the fuel load would
increase. Today, species of Baeckia, Kunzea and
Leptospemum are dominant on all of the swamps
(Chalson and Martin, this volume). There is a trend
for Leptospermum and Kunzea species to increase
slightly in the late Holocene, with a further increase
in the European Zone. These woody species would
have had the capacity to produce more charcoal when
burnt and be incorporated in the sediments, especially
when growing on the swamp, when compared with
the smaller sedges and reeds.
At Mountain Lagoon (Robbie and Martin, 2007),
fire activity was low through the Holocene, until about
3-2 ka, when it increased. This pattern is similar to
those of this study.
At Gooches Crater the charcoal content and
hence fire activity fluctuates between 14 ka and 9
ka, then follows a period of low fire activity until
about 6 ka, then a period of dramatic increase in fire
activity in the late Holocene (Black and Mooney,
2006). Fire activity reaches unprecedented levels in
the post-European period (Black and Mooney, 2006).
The increase in fire activity in the mid Holocene
is attributed to climate, in particular to the greater
seasonality associated with the onset of the El Nifio-
Proc. Linn. Soc. N.S.W., 130, 2009
Southern Oscillation (ENSO) phenomenon (Black et
al., 2007). This pattern of fire activity is similar to that
seen in the sites of this study.
At Kings Waterhole Swamp (Black and Mooney,
2007), the fire activity was low about 6 ka, then
mecreased between 5-3 ka, after which it decreased to
low levels to the present. This pattern of fire activity
is quite unlike that of this study. It is thought that
the decline in fire activity after 3 ka represented
an alteration to Aboriginal management strategies
associated with increasing population and/or the
increased risk of conflagration in an ENSO-dominated
climate (Black and Mooney, 2007).
Black et al. (2007) examined the charcoal record
together with the archaeological record in an attempt
to assess the likely effect of Aboriginal burning on the
ecosystem. At Gooches Crater Swamp, the charcoal
content appeared to be most influenced by climate,
with an abrupt increase in the mid Holocene, perhaps
associated with the onset of the modern ENSO-
dominated conditions. Kings Waterhole also showed
the abrupt increase in the mid Holocene, but there was
a marked decrease in charcoal from about 3 ka. Lake
Baraba also showed similar low levels of charcoal in
the late Holocene. The archaeological records of all
three regions showed increased activity/habitation in
the late Holocene. It is thus possible that Aborigines
strongly influenced fire activity in some places in the
Sydney Basin during the late Holocene to prevent the
risk of large intense fires as the ENSO-dominated
climate became more prevalent (Black et al., 2007)
CONCLUSIONS
Seven swamps were studied and each had its
own distinctive history. Where the Eucalyptus species
were identified, the dominant species were different
at each site, as they are today.
Similarities in the histories could be seen between
some of the sites and are as follows:
In the early Holocene, the vegetation was more
wooded, i.e. woodland or forest, which suggests a
warmer wetter climate. Only the three sites at the
highest altitudes had sediments of early Holocene
age.
In the mid Holocene, the vegetation was less
wooded in the three highest sites when the vegetation
was probably shrublands and heaths, and this suggests
a drier climate. Other sites at lower elevations were
wooded in the mid Holocene.
The Eucalyptus species return to the less
wooded sites towards the late Holocene. There is
also a tendency for an increase in Baeckea, Kunzea,
105
HOLOCENE HISTORY OF BLUE MOUNTAINS VEGETATION
Leptospermum and Melaleuca, the woody shrubs.
These woody shrubs are dominant on the swamps
today.
There is some decline in Casuarinaceae in the
European period but the Eucalyptus species are
maintained at about the same level as in the late
Holocene. The woody swamp shrubs increase in the
late Holocene and European period.
The charcoal levels suggest that there was moderate
fire activity in the early Holocene when the vegetation
was more wooded, decreased fire in the mid Holocene
when the vegetation was more open, with increased
fire in the late Holocene and a further increase in the
European period.
It is thought that the altered fire regime under
European settlement encouraged the increase in
woody shrubs on the swamps (and elsewhere) which
in turn produced more charcoal.
These swamps on sandstone are highly erodable
and a sand body at about 1.2-1.6 ka in the three most
northerly and easterly swamps suggests they may
have suffered an erosive event about that time. The
destabilising event(s) which triggered this erosion is
uncertain.
ACKNOWLEDGEMENTS
We are indebted to the Joyce W. Vickery Research
Fund of the Linnean Society of New South Wales, 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, Dr. Mike Barbetti, 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
Allen, R. and Lindesay, J. (1998). Past climates of
Australasia. In: ‘Climates of the Southern Continents’
(Eds J.E. Hobbs, J.A. Lindesay, H.A. Bridgman) pp.
208-247. (Wiley & Sons, Chichester).
Attenbrow, V. (2002). “‘Sydney’s Aboriginal Past’.
(University of New South Wales Press. Sydney) 225
pp.
Australian Government, Department of the Environment
and Water Resources (2007a). World Heritage,
Blue Mountains. (http://www.environment.gov.au/
worldheritage/sites/blue/index/html) Accessed May,
2007.
106
Australian Government, Department of the Environment
and Water Resources (2007b). Temperate highland
peat swamps on sandstone. (http://environment. gov.
au/biodiversity/threatened/communities/temperate-
highland-peat-swamps.html). Accessed October
2007).
Bembrick, C. (1980). Geology of the Blue Mountains,
western Sydney Basin. In “A Guide to the Sydney
Basin’ Eds C. Herbert and R. Helby, pp 135-161
(Geological Survey of Sydney NSW Bulletin No. 26,
Sydney).
Bembrick, C., Herbert, C, Scheibner, E. and Stuntz, J.
(1980). 1. Structural subdivision of the Sydney Basin.
In “A Guide to the Sydney Basin’ Eds C. Herbert and
R. Helby, pp 3-9 (Geological Survey of Sydney NSW
Bulletin No. 26, Sydney).
Birks, H.J.B. and Birks, H.H. (1980) ‘Quaternary
Palaeoecology’. (Edward Arnold, London).
Black, M.P. and Mooney, S.D. (2006). Holocene fire
history from the Greater Blue Mountains World
Heritage Area, New South Wales, Australia:
the climate, human and fire nexus. Regional
Environmental Change 6, 41-51.
Black, M.P. and Mooney, S.D. (2007). The response of
Aboriginal burning practices to population levels
and El Nifio-Southern Oscillation Events during the
mid- to late-Holocene: a case study from the Sydney
Basin using charcoal and pollen analysis. Australian
Geographer 38, 37-52.
Black, M.P., Mooney, S.D. and Haberle, S.G. (2007). The
fire, human and climate nexus in the Sydney Basin,
eastern Australia. The Holocene 17, 469-480.
Black, M.P., Mooney, S.D. and Martin, H.A. (2006). A >
43,000-year vegetation and fire history from Lake
Baraba, New South Wales, Australia. Quaternary
Science Reviews 25, 3003-3016.
Boland, D.J., Brooker, M.I.H., Chippendale, G.M., Hall,
N. H., Hyland, B.P.M., et al. (2002). ‘Forest Trees
of Australia’ 4" Edition. (Thomas Nelson: Australia)
687 pp.
BoM, (2006). Commonwealth Bureau of Meteorology
Website (http://www.bom.gov.au). Accessed August
2006.
Breckell, M. (1993). Shades of grey: early contact in the
Blue Mountains. In “Blue Mountains Dreaming, the
Aboriginal Heritage’ (Ed. E. Stockton) pp. 114-121
(Three Sisters Publication Winmalee NSW).
Brown, C.A. (1960). ‘Palynological Techniques’ (C.A.
Brown, Baton Rouge) 188 pp.
Bureau of Meteorology (1979). ‘Climatic survey Sydney
Region 5 New South Wales’. Depatment of Science
and the Environment, Commonwealth of Australia
(Australian Government Publishing Service,
Canberra) 141 pp.
CalPal (Version March 2007). Cologne Radiocarbon
Calibration and Palaeoclimatic Research Package.
(http://www.calpal.de/). Accessed May 2008.
Chalson, J.M. (1991). The late Quaternary vegetation
and climatic history of the Blue Mountains, NSW,
Proc. Linn. Soc. N.S.W., 130, 2009
J.M. CHALSON AND H.A. MARTIN
Australia. PhD Thesis, University of New South
Wales (unpubl.)
Chalson, J.M. and Martin, H.A. (1995). The pollen
morphology of some co-occurring species of
the family Myrtaceae from the Sydney region.
Proceedings of the Linnean Society of New South
Wales 115, 163-191.
Chalson, J.M. and Martin, H.A. (2008). A 38,000 year
history of the vegetation at Penrith Lakes, New South
Wales. Proceedings of the Linnean Society of New
South Wales 129, 97-111.
Chalson, J.M. and Martin, H.A. (this volume). Modern
pollen deposition under vegetation of the Blue
Mountains, New South Wales. Proceedings of the
Linnean Society of New South Wales 130, 111-137.
Donders, T.H., Haberle, S.G, Hope, G., Wagnera, F.
and Visschera, H. (2007). Pollen evidence for the
transition of the Eastern Australian Climate system
from the post-glacial to the present-day ENSO mode.
Quaternary Science Reviews 26, 1621-1637.
Entwisle, T. (2005). She-oak up in smoke. Nature
Australia, Spring 2005 28(6), 72-73.
Keith, D.A. and Benson, D.H. (1988) The natural
vegetation of the Katoomba 1:100 000 map sheet.
Cunninghamia 2, 107-143.
Kohen, J. (1995). ‘Aboriginal Environmental Impacts’.
(University of New South Wales Press, Sydney) 160
pp.
Langford-Smith, T. (1979). Geomorphology. In “An
Outline of the Geology and Geomorphology of the
Sydney Basin’ (Ed. D. Branagan), pp 49-58. (Science
Press, Marrickville).
Moore, P.D., Webb, J.A. and Collison, M.E. (1991).
“Pollen Analysis’. (Blackwell Scientific Publications,
Oxford) 216 pp.
Moy, C.M.., Seltzer, G.O., Rodbell, D.T. and Anderson,
D.M. (2002). Variability of El-Nifio/Southern
Oscillation activity at millennial timescales during
the Holocene epoch. Nature, 420, 162-165.
Robbie, A. and Martin, H.A. (2007). The history of
the vegetation from the last glacial maximum at
Mountain Lagoon, Blue Mountains, New South
Wales. Proceedings of the Linnean Society of New
South Wales 128, 57-80.
Rose, S. and Martin, H.A. (2007). The vegetation history
of the Holocene at Dry Lake, Thirlemere, New South
Wales. Proceedings of the Linnean Society of New
South Wales 128, 15-55.
Sainty, G.R. and Jacobs, S.W.L. (1981). “Water Plants of
New South Wales’ (Water Resources Commission
New South Wales, Sydney) 550 pp.
Smith, J. (1993). Katoomba’s fringe dwellers. In “Blue
Mountains Dreaming, the Aboriginal Heritage’ (Ed.
E. Stockton) pp. 122-135. (Three Sisters Publication
Winmalee NSW).
Stockton, E. (1993a). Archeology of the Blue Mountains.
In ‘Blue Mountains Dreaming, the Aboriginal
Heritage’ (Ed. E. Stockton) pp. 23-54 (Three Sisters
Publication Winmalee NSW).
Proc. Linn. Soc. N.S.W., 130, 2009
Stockton, E (1993b). Aboriginal sites in the Blue
Mountains. In “Blue Mountains Dreaming, the
Aboriginal Heritage’ (Ed. E. Stockton) pp. 55-62
(Three Sisters Publication Winmalee NSW).
Sullivan, L. (2007). Small victory in long battle to save
hanging swamps. Sydney Morning Herald Letters, 17
October 2007.
Ward. D.J., Lamont, B.B. and Burrows, C.L. (2001).
Grasstrees reveal contrasting fire regimes in
eucalypt forest before and after European settlement
of southwestern Australia. Forest Ecology and
Management 150, 323-329.
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
SEMEN RUA
fel ath Ce lt eal
saath mst dagen sid maces st we en
— Drench
3 eee castes iat Lise ap cata
nt ne “ome vat Oli: a hich ELeTIIS es 1 ? Siac ater
ms vs ; y sented hordes
cle 4 tt a or alge
on i a | : ; piper ae eine wi ied hil eer o.
1
LOOTED ) OF whee AW erty pw be er anal
aoe yet NM Yrb als 2 RPA DAH LE asic istry” ae ala honk “3
einoge terns Sele OT ieee {Pec mahiy’ PHP os ND athe SCC | ys
oreo tt rite ybose beta inesttiay' foeen ey an ea i el ae nih eens
abiue qaob te: eeny Vente ib Wh ainsty ag. a ocak wale preven
line skiirrsiiny Huby Senad TOWwG ERAS Tat eae tS ehh FT ete a9) ew sat! hee sear ETH iy au
as ahiow' vbnat ,chirtstet yi PRS ty erred H gigas Hoe mg
"WHO? viotiae 76 [ndekaale 1d ES hy Ivins i ww” me hams de a
tee Baie tay ene Re iit fear “rh thy vorank 208. Res, Axprallyy sN
Savi y weatho obit. vloneist bili Wisk tbh Wy eT eerie Kare vee
ic vest: SEM MR a i ihe Pies ey a
ation olin mi pst Hey a eee he leas er na os.
| hc ( uke mingle ay
ot eH hrasatnih 5 2 ea ania
(FOR GREY sere exgy! ANG PUERRDN Ay ASH Se Co nas aati
slid, "Snake jonah diy eboney tt Finmeat eq eet
Cneviltedaphd): scleriphy dans uinkeuorey® M anh) savienihe
fest onetlbotaye 1h yuh ee RB cel OW SRY mihi eLGe wonalnls a ani
mE xoteaniinsh Pedi bias Ai ea Su ifobebe! J
\ Wf i Were 4 vee gaclor
liye dabg espii dO, tiaslaabee, bp, thsad LOTUS 0 ae
eee Alot TAR AHURA DITA , aa WE?
AWE. brs estrada ONG Lao a> sey BA ERS Pres he “SStlolisiye pSaalhiwt
| x} mati THRE Hite OMS ano aulic salaly | ahiohlaneige
Hy 1 SP): MTCTO Pay thous anisherstg Gl wach isis
si ry’ a eel gahay age Mm Llale oh te) Bae
{ (Oe T pneielog > 598 oa nolan, i RITA L 2) Maal st er D940 ounle
waeyyy ( eh ep att tO He bia Be
ty Vide tht fred ot he [ev (ts TT “alg fv :
(fRO4) mosien) ou? gp Sa RANGA? i oh Cl oqyt sad
poe aidedisaHhy Hoet'ita ss Be eee 0d aaa seat ani) i
me, incarceration iarnnes op ee ye balhe — sana
rah! SEP RA T PiObahity % inka how bere,
pataltiy Qader ics aprpiainlanen ach neha thee rh vt ge
Saath hn sity ane ery ye! eneaptey} taht ar. wANC Ls waterrle Shute ; ,, =
- ; ~
ee
‘parotid bresrts, dss cat deri. growl aren Ww mor! i
ert ene Lwutifionas a wy ray et winnd SN U “one wubrtie nily Livihiiarae tebe pe
1 q 7 r
Dy. ico ip ' a. THB siwns fi¢ho nate yes ee, ti aa 1 he sear titite. } nbudifion he! snaibeland
4 Piberpa ag aia ehcp oder égald tobmedit: ” ane: Ns
ead pad, nie eer Sarwan — a oti A hae tn Here _ a cer ha ll A ee
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
\ oy be L Grose ®
pleekheath
bos
Mediow Bath &
Study area
ney
Burralow
k Sw. *
oe eee
~A
y, S
=i Springwood & ie
‘ Katoomba | awson sj Qa 2 Penrith
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
eo] @ eu]
a5 Ro tty
oe? § é g
Ss % 0
& G @ & & @
Se g = @ a
oY s y fo} Lg &
C) oy 2 & @ a a fr io)
72 oP g § 2 e e 8g &
&F § & é g § <« << 2
2 IY YL te) RS ae :
N SoS & & J é eo é é
on ee ek : : eves ©
Burralow Creek Swamp
1 Mid swamp S —
2 Mid swamp ,
3 Swamp edge
4 Weadiand | eee oe
10 Mid swamp t |
11 Swamp edge
12 Mid swamp z |
13 Swamp edge
18 Woodland
16 Mid swamp iz L
Swamp edge a
21 Forest
os
a m7
N fh
+ HW
Swamp edge
26 Forest ae:
Mid swamp |
Swamp edge {
Woodland |
mh
o oo ~
6 Mid swamp fal
7 Midswamp [~ ca
8 Swamp edge
9 Woodland ia ~~ a
Berea | direc tee
Warrimoo Oval Swamp
Notts Swamp
—
Ingar Swamp
| eee
Kings Tableland Swamp
fay oF ATTA, MaapepD
er mee sae
RES ie, ae
Rye oe Ge
i
I
Katoomba Swamp
Toles Bel al dele TT jaar ase - th,
ay te ee
Newnes Swamp
% ee |
| aos Scale —— ee count a
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
Ca ve rd % & ® $ S ¢3 F) £
~ 4 @ &
So 2 8b 9 FE EE SSE GET ES
®2¢ F S&S Fs SF EFESESES FEA BEEE
OL LIF EES LETC LLEP TIE
Burralow Creek Swamp ab
7 Mid swamp | | | i L j } ! '
2 Mid swamp | | is \ | r
3 Swamp edge |
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
s ¢ e
S) §
oe
© S &
~ op 5 = @
g &$ s &8 & £ Fe
® & © § £ v § gs
Lo §& & §€ & S S RY
Q + S FF LN WG > & a) > §
eé ree & & ? g P #8
ff etd & 3 i ee Be
\ ‘ ‘ | aoe! th aati
Woodland 10ar
Woodland 10ffita
29 i
30 | |
31 jae
32 |
Open Woodland 11a
39 ae
AE cial Saal Be Ao 3
Open forest 10ag
5 -——
at | | [
pf
ier es
= |I| |E
a
Ss
Open forest 9i
Tall Open Forest 6c
{0a ae
Open Forest 10f
= ARE E| res
F
7
ic}
3
3
Open Heath 2%f, 21d, 21c
22 Scale a
33 pe fe i rier =
34 20% of total count j
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
€
§ og
v * Scale 1, 20% of total count Bo
Si S £&
cS S $s
< & &¢ @
Be anc ghesirescom Sle
o ae 7 % oe ¢
Sea askS ao G @ © o §
2LLESS ESa SF EBES F §
SESS EE FFE S 5 LL oe & & §
ELLES SF EEESESS FEE
Bes eiatate te ieakodik ut fo an
eee t Day => | ieee take | ae
Woodland 10ar
|
1
at fr Ha se Wa Ba | |
14 | ela : | | ; | |
15° Bhi i r r
19 | | | | lest t F F
Woodland 10f/i1a
29 [> Be We
SOC Pelee ce
31 r | LF Poy t Ravel
gb hie PLE A re La
Open Woodland 11a
Sat tiiie fe Abe seprte Hey nies oF
sn . | arse aa cr
Open forest 10ag
| | — Pot
|. \ ie
i la. doal
er | ror
| L nen, hee Cake eee ace er ws
AE stad - |
ares ; - +
Herald |
i it ones ee
Oo socaenpeccor
i
=
o
®
m
2
| Open Forest 6c
ill
ee
pen Forest 10f
Or ae ee ee | mF
io sorrel sal psligee aaa ie
Open Heath 21f, 21d, 21c
iat. fk
TER PE y+]
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 =| &
Oo mo oa RN
aie te. fe ee eee eet
; a
oy es ea Tt
wae ee |
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
REFERENCES
Amsden, T. W. (1968). Articulate brachiopods of the St. Clair
Limestone (Silurian), Arkansas, and the Clarita
Formation (Silurian), Oklahoma. Paleontological
Society Memoir 1.
Barrande, J. (1847). Uber die Brachiopoden der silurischen
Schichten von Bohmen. Naturwissenschaftlichen
Abhandlungen. I Band. W. Haidinger, Wien, 357-
475. [not seen, fide Havliéek, 1982].
Cooper, G.A. (1956). Chazyan and related brachiopods.
Smithsonian Miscellaneous Collections 127.
Hall, J, and Clarke, J.M. (1893). An introduction to the
study of the genera of Palaeozoic Brachiopoda.
Palaeontology of New York 8 (2), 1-317.
Havli¢ek, V. (1982). New genera of rhynchonellid and
camerellid brachiopods in the Silurian of Bohemia.
Véstnik Ustredniho tistavu geologického 57, 365-
372.
Havlicek, V. (1992). New Lower Devonian (Lochkovian-
Zlichovian) rhynchonellid brachiopods in the Prague
Basin. Sbornik Geologickych Véd, Paleontologia
32, 55-122.
Havliéek, V. and Storch, P. (1990). Silurian brachiopods
and benthic communities in the Prague Basin
(Czechoslovakia). Rozpravy Ustredniho Ustavu
Geologického 48.
Jenkins, C. (1879). On the geology of Yass Plains.
Proceedings of the Linnean Society of New South
Wales, series 1, 3, 21-32.
Kuhn, O. (1949). ‘Lehrbuch der Palaozoologie’. (E.
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
ordovika 1 silura Tuvy. Trudy Institut Geologii i
Geofiziki, Sibirskoye otdeleniye Akademiya Nauk
SSSR 635. [Russian]
Laurie, J.R. (1991). Articulate brachiopods from the
Ordovician and Lower Silurian of Tasmania. Memoir
of the Association of Australasian Palaeontologists
11, 1-106.
Mitchell, J. (1887). On some new trilobites from Bowning,
N.S.W. Proceedings of the Linnean Society of New
South Wales, series 2,2, 435-440.
Plodowski, G. (1973). Revision der Brachiopoden-Fauna
des Ober-Siluriums der Karnischen Alpen, 2.
Rhynchonellacea aus den Zentralkarnischen Alpen.
Senckenbergiana lethaea 54, 65-103.
Rong J.-Y., Boucot, A.J., Su, Y.-Z. and Strusz, D.L.,1995.
Biogeographical analysis of Late Silurian
brachiopod faunas, chiefly from Asia and Australia.
Lethaia 28, 39-60.
Savage, N.M., Mancefiido, M.O., Owen, E.F., Carlson,
S.J., Grant, R.E., Dagys, A.S. and Sun D.-L.,
2002. Rhynchonellida. In ‘Treatise on Invertebrate
Paleontology, Part HH, Brachiopoda, revised,
volume 4: Rhynchonelliformea (part)’ (Ed. R.L.
Kaesler), pp. 1027-1376. (Geological Society of
America, Denver and University of Kansas Press,
Lawrence).
Schmidt, H. (1965). Neue Befunde an Paldozoischen
Rhynchonellacea (Brachiopoda). Senckenbergiana
Lethaea 46, 1-25.
Schuchert, C. (1913). Class 2. Brachiopoda. In ‘Text-
book of Palaeontology, volume 1, part 1, edition 2,
translated and edited by Charles R. Eastman’ (Ed.
K.A. von Zittel) pp. 355-420. (MacMillan and Co.,
Ltd, London).
Shearsby, A.J. (1912). The geology of the Yass district.
Reports of the 13" Meeting of the Australasian
Association for the Advancement of Science, Sydney
1911, 106-119.
Strusz, D.L. (2002). Brachiopods of the Orders Protorthida
and Orthida from the Silurian of the Yass Syncline,
southern New South Wales. Alcheringa 26, 49-86.
Strusz, D.L. (2003). Late Silurian strophomenate
brachiopods from Yass, New South Wales.
Alcheringa 27, 1-35.
Strusz, D.L. (2005). Late Silurian pentameride brachiopods
from Yass and Molong, New South Wales.
Alcheringa 29, 205-228.
Williams, A. and Brunton, C.H.C. (1997). Morphological
and anatomical terms applied to brachiopods. In
“Treatise on Invertebrate Paleontology, Part H,
Brachiopoda, revised, volume 1: Introduction’ (Ed.
R.L. Kaesler) pp. 423-440. (Geological Society of
America, Boulder, and University of Kansas Press,
Lawrence).
Williams, A., Carlson, S.J., Brunton, H.C., Holmer, L.E. and
Popov, L. (1996). A supra-ordinal classification of
the Brachiopoda. Philosophical Transactions of the
Royal Society, London, Series B, 351, 1171-1193.
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
REFERENCES
Bougher, N. L. & K. Syme, (1988). ‘Fungi of Southern
Australia’. (University of Western Australia Press,
Perth ).
Brandrud, T. E. (1983). Cortinarius subgen. Cortinarius
(Agaricales) in the Nordic countries, taxonomy,
ecology and chorology - Nordic Journal of Botany
3, 577-592.
Brandrud, T. E. et. al (1989-1998) Cortinarius. Flora
Photographica. (Cortinarius HB, Matfors, Sweden).
Breitenbach, J. &F. Kranzlin, (2000). Fungi
of Switzerland. Vol 5. Cortinariaceae.
(Lucerne,Switzerland).
Brittlebank, C. C. (1940). Catalogue of Australian Fungi.
(Published by the author )
Chambers, S. M. & N. A. Sawyer & J.W.G. Cairney
(1999). Molecular identification of co-occurring
Cortinarius and Dermocybe species from
southeastern Australian sclerophyll forests.
Mycorrhiza 9, 85-90.
Cleland, J. B. (1934). ‘Toadstools and -Mushrooms
and Other Larger Fungi of South Australia’. 1.
(Government Printer, Adelaide).
Cooke, M. C. (1892). “Handbook of Australian Fungi’.
(Williams & Norgate, London.)
Garnica, S., Weiss, M., Oertel, B., Oberwinkler, F.,
(2005). A framework for phylogenetic classification
in the genus Cortinarius (Basidiomycota, Agricales)
derived from morphological and molecular data.
— Canadian Journal of Botany 83 (11), 1457 — 1477.
Gasparini, B. (2001). A Contribution to the knowledge of
Cortinarius and allied genera of Southern Tasmania,
Australia. 1. Cortinarius subgenus Cortinarius.
Australasian_Mycologist 20 (1), 49-54.
Grgurinovic, C. A. (1997). “Larger Fungi of South
Australia’. (Botanic Gardens and State Herbarium
and Flora and Fauna of South Australia Handbooks
Committee, Adelaide).
Griffiths, K. (1985). ‘A Field Guide to the Larger Fungi
of the Darling Scarp and the South West of Western
Australia’. (Published by the author).
Hilton, R. N. (1983). A census of the larger fungi of
Western Australia. Part 11 — Journal of the Royal
Society of Western Australia 70, 111-118.
Hoiland, K. & A. Holst-Jensen, (2000). Cortinarius
phylogeny and possible taxonomic implications of
ITS rDNA sequences. - Mycologia 92(4), 694-710.
Horak, E. (1979). Fungi, Basidiomycetes Agaricales y
Gasteromycetes Secotioides. Flora Criptogamica de
Tierra del Fuego. Tomo X1 Fascicle 6 (Buenos Aires,
Argentina).
Horak, E. (2005). ‘ Rohrlinge und Blatterpilze in Europa.’
(Elsevier, Munchen).
Knudsen, H. & Vesterholt, J. (Eds).(2008). “Funga
Nordica’. (Nordsvamp, Copenhagen).
Kornerup, A. & Wanscher, J. H. (1978). ‘Methuen
Handbook of Colour’. (Methuen, London).
Proc. Linn. Soc. N.S.W., 130, 2009
Maerz, A. & M. R. Paul (1950) “A Dictionary of Colour’.
2nd Edition. McGraw —Hill, New York.
May, T. W. & A. E. Wood, (1997) Catalogue and
Bibliography of Australian Macrofungi. 1.
Basidiomycetes p.p. Fungi of Australia Volume 2A.
Australian Biological Resources Study. Canberra.
348 pp.
McAlpine, D. (1895). Systematic Arrangement of
Australian Fungi, (R. S. Bain, Government Printer,
Melbourne).
Moser, M. (1983). Keys to Agarics and Boleti
(Polyporales, Boletales, Agaricales, Russules)
(Roger Phillips. London).
Moser, M. M. (1986) 1987. Cortinarius Fr. subgenus
Cortinarius 1n the SW-Pacific area. Sydowia 39 :
138-147
Moser, M. M. & E. Horak, (1975). Cortinarius Fr. und
nahe verwandte Gattungen in Sudamerika, Beih.
Nova Hedwigia 52: 1-628.
Shepherd, C. J. & C. J. Totterdill (1988). Mushrooms and
Toadstools of Australia. (Inkata Press. Melbourne).
Syme, K. (1992) Survey of the Larger Fungi of the Two
Peoples Bay Nature Reserve. Denmark Environment
Centre, Denmark.
Young, A. M. (1994) Common Australian Fungi. (New
South Wales University Press, Sydney).
155
_ ConTINARIL AY PF: sti RC vamewcnten USINA AUS
Sool a seein “OHO Tle 5 Ds Aystnaliat
Ato? wot DH winidoM cvtiba pe
. baw sugolmis's {FOP pa bos OF Da, Be ET Yauch
[ender A onl YeuAdo AeptergoildiG |
AS oat attecA to Liat |e strpvirnal Met
ensian) obi vsirabio8 Bo pia i neifeugerd
Pa } Aa) phe
w re | durgotet} URL) ) tein) AIM
tahoe T smears viata, ane em east i mellow:
ene \
isto bai xaieguscn htor pee M, rsen
dicvitenid A colar saab on
{ovaling,S ml i
Ht hte Ai dvdr’ wens’ hie
OF ive’. .eoam we aKwe av or i ni 74
ine 24 zaioniiws. lastinlbars Le eae
Ais8 cchnatiot ai cogmmtisd otbaew tee olen
Lime ie raiaiadiA erg
tiem.oninrteuy (eet) f As
(umiadie wean vee tat) pon Ton choot .
newt dy lot won adit to fev (tet Rw do
iTramowena: Panel! pirat oR eroaei dn asta}
drmerstoKh a he
wavy gel wnnaleendeae onion (Doe MA
{ vonbu® bayr'l 4 NAB Frist Uae frend
| } |
fry oy
} j j
; / \ ;
a \ |
, |
@y. \ 1}
AJ \
ri \ }
( "> C4
} f
Vapeneh a Contieandts hat Lrwellvresia (PERTH acai ETO ae Billed A PAE
d.. (ear yiiie
rita abumient, drow filer of
ae
Heal caticle a teat,
pearcels
iveted layer SUI) ate deen, ..compored
SUT ALS
apemial
i eet <
aT at LY Orne ie
eie, Ane
a+ 4 y
ittenwed, Dat
meter, thst
witly.no
}
Mme. Cysin
Ho one Wer the
Onova amore denacly
i Hh CSHSCS On PHik bly IT Siporss
eid eiga?
is (et oe ae
rary ie aundattani
3 bwial fener wiht Nas
or whether 1 waa
iyibeaa Mana woth, Wie eaatic art ria isceur
hice ce eeerve adlectmng ony
We enwintend,
j : ee
PULTE OT PILES Vrs
ce Reeder tis) ST ar 3S
paneer) te the wintebed hoe
eperitem:
ary Thigy Meh
ert? PER Cee art
mot j
iu callovtven), Ti Qe pore oF
anhselien at:
has
inf
‘ynvdnor fs
“aban eh
* | j , 7 ae
evil nlperey > (NS
- habbon ental .
tghas A en i, sisal
} dae oti % 1)
tS ia a a ried tote)
Be Anite rae. wey
as agi
Soc ak styiatalod, amine vat
as-Gatiow) wu
pre ee pane
ait DS rane paisa sf
dihalben-+-R—rigaloy AA, sinh
diaeie Yo igre yoyiat” ATOR TA 9 orvunh
nushedtaH ssid bie anobwO ey ‘ellen
tye rie i igh in sbivid bal q re Aeber) Ae
. ‘one ‘
cri) sb ie —" pie fe rite
isiy 3 re “OT theres bh rvored YG Gk
ekvenie> (0000 )\c5e0nl-elol AD A De
Jo: RnodnoGM 2imonoxs! aidigang bas ynogotydle
ODO: (MAE ONRIE Didi tol RCs. ani
| Yableabiag Avatzoymnnibiesdl inn en
ob Bharani Derek mahypetowe’ os
Nene Meet} «) oigtaeetls iK ov ont
ey UA ee ee oe Mew South phason ,
wf iQirindtiei Svbeepaniatid Bewerpnilade be 48
“tor penmnisiith lo motlee saciimcteottog M
control: Maou LBOOGjx dead wl § nt Wks ie
lot axes: imenforgudasgod epoca eal ay
nautitioM’ (ATOL) Hi, ger ie
adbio.t — Sag le ta
Wi,
phat gun
(PE
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.
REFERENCES
Bancroft, B.B. (1933). Correlation tables of the stages
Costonian-Onnian in England and Wales. 4pp.
Printed and published by the author, Blakeney,
Glouchestershire, England (not sighted; fide Williams
et al. 2000).
Carlson, S.J. (2002). Suborder Syntrophiidina. In ‘Treatise
on Invertebrate Paleontology, Part H. Brachiopoda
(Revised), Vol. 4’ (Eds A. Williams, C.H.C. Brunton
and S.J. Carlson) pp. 929-960. (The Geological
Society of America: Boulder, and The University of
Kansas Press: Lawrence).
Cocks, L.R.M. (2005). Strophomenate brachiopods
from the Late Ordovician Boda Limestone of
Sweden: their systematics and implications
for palaeogeography. Journal of Systematic
Palaeontology 3, 243-282.
Cocks, L.R.M. and Rong, J.-y. (2000). Order
Strophomenida. In “Treatise on Invertebrate
Paleontology, Part H. Brachiopoda (Revised), Vol. 2’
(Eds A. Williams, C.H.C. Brunton and S.J. Carlson)
pp. 216-348. (The Geological Society of America:
Boulder, and The University of Kansas Press:
Lawrence).
Cocks, L.R.M. and Zhan, R.-b. (1998). Caradoc
brachiopods from the Shan States, Burma
(Myanmar). Bulletin of the Natural History Museum,
London (Geology) 54, 109-130.
Cooper, G.A. (1956). Chazyan and related brachiopods.
Smithsonian Miscellaneous Collections 127, 1245 pp.
+ 269 pl.
Edgecombe, G.D. and Webby, B.D. (2006). The
Ordovician encrinurid trilobite Sinocybele from New
176
South Wales and its biogeographic significance.
Memoirs of the Association of Australasian
Palaeontologists 32, 413-422.
Edgecombe, G.D. and Webby, B.D. (2007). Ordovician
trilobites with eastern Gondwanan affinities from
central-west New South Wales and Tasmania.
Memoirs of the Association of Australasian
Palaeontologists 34, 255-281.
Klenina, L.N. (1984). Brakhiopody 1 biostratigrafiya
srednego 1 verkhnego ordovika Chingiz-
Tarbagatayskogo megantiklinortya. [Brachiopods and
biostratigraphy of the Middle and Upper Ordovician
of the Chingiz-Tarbagatai Meganticlorium].
In Klenina, L.N., Nikitin, I.F. and Popov, L.E.
‘Brakhiopody 1 biostratigrafiya srednego 1 verkhnego
ordovika khrebta Chingiz’, pp. 6-125. (Nauka, Alma-
Ata).
Laurie, J.R. (1991). Articulate brachiopods from the
Ordovician and Lower Silurian of Tasmania. In
P.A. Jell (ed.) “Australian Ordovician Brachiopod
Studies’. Memoirs of the Association of Australasian
Palaeontologists 11, 1-106.
McLean, R.A. (1974). The geology of the Regans
Creek area, near Cargo, central New South Wales.
Proceedings of the Linnean Society of New South
Wales 98, 196-221.
McLean, R.A. and Webby, B.D. (1976). Upper Ordovician
rugose corals of central New South Wales.
Proceedings of the Linnean Society of New South
Wales 100, 231-244.
Neumann, R.B. (1994). Late Ordovician (Ashgill)
Foliomena fauna brachiopods from northeastern
Maine. Journal of Paleontology 68, 1218-1234.
Nikiforova, O.I. and Andreeva, O.N. (1961).
Biostratigrafiya Paleozoya Sibirskoi Platformi, vol.
|. Stratigrafiia Ordovika 1 Silura Sibirskoi Platformy
i ee Paleontologichesko Obosnovanie (Brakhiopody)
[Biostratigraphy of the Paleozoic of the Siberian
Platform, vol. 1. Ordovician and Silurian stratigraphy
of the Siberian Platform and its paleontological
basis (Brachiopoda)]. Vsesoiuznyi Nauchno-
Issledovatel skii Geologicheskii Institut (VSEGED),
Trudy 56, 1-412.
Nikiforova, O.I. and Sapelnikoy, V-.P. (1973). Nekotorye
dreynie pentameridy Zeravshanksogo khrebta.
[Some fossil pentamerids from the Zeravshan
Range]. In Sapelnikov, V.P. and Chuvashoy, B.I.
(eds), Materialy po paleontologii srednego Paleozoia
Uralo — Tian’shan’skoi oblasti. Akademiya Nauk
SSSR, Ural ’ski Nauchnyi Tsentr, Institut Geologii I
Geokimii, Trudy 99, 64-81.
Nikitin, I.F. and Popov, L.E. (1983). Sredneordovikskie
ortatsei 1 plektambonitatsei severnogo Priishimya 1
basseyna reki Akhzhar v Tsentral’nom Kazakhstane
(Brakhiopody). [Middle Ordovician orthids and
plectambonitaceans from northern Priishimiya
and the River Azkhar in central Kazakhstan
(Brachiopoda)]. Ezhegodnik Vsesoyuznogo
Paleontologicheskogo Obshchestva 26, 228-247.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Nikitin, I.F. and Popov, L.E. (1984). Brakhiopody
bestamakskoy i sargaldakskoy svit (srednii ordovik).
[Brachiopods from the Bestamak and Sargaldak
Formations (Middle Ordovician)]. In Klenina,
L.N., Nikitin, I.F., and Popov, L.E. “Brakhiopody
i biostratigrafiya srednego 1 verkhnego ordovika
khrebta Chingiz’, pp. 126-166. (Nauka, Alma-Ata).
Nikitin, I.F. and Popoy, L.E. (1996). Strophomenid and
triplesiid brachiopods from an Upper Ordovician
carbonate mound in central Kazakhstan. A/cheringa
20, 1-20.
Nikitin, I.F., Popov, L.E. and Bassett, M.G. (2006).
Late Ordovician rhynchonelliformean brachiopods
of north-eastern Central Kazakhstan. In Bassett,
M.G. and Deisler, V.K. (eds) “Studies in Palaeozoic
Palaeontology’. National Museum of Wales
Geological Series No. 25, pp. 223-294. (National
Museum of Wales, Cardiff).
Nikitin, I.F., Popov, L.E., and Holmer, L.E. (1996). Late
Ordovician brachiopod assemblage of Hiberno-
Salairian type from Central Kazakhstan. Geologiska
Foéreningens i Stockholm Férhandlingar 118, 83-96.
Percival, I.G. (1976). The geology of the Licking Hole
Creek area, near Walli, central western New South
Wales. Journal and Proceedings of the Royal Society
of New South Wales 109, 7-23.
Percival, I.G. (1978). Inarticulate brachiopods from the
Late Ordovician of New South Wales, and their
palaeoecological significance. A/cheringa 2, 117-141.
Percival, I.G. (1979a). Ordovician plectambonitacean
brachiopods from New South Wales. A/cheringa 3,
91-116.
Percival, I.G. (1979b). Late Ordovician articulate
brachiopods from Gunningbland, central western
New South Wales. Proceedings of the Linnean
Society of New South Wales 103, 175-187.
Percival, I.G. (1991). Late Ordovician articulate
brachiopods from central New South Wales. In
P.A. Jell (ed.) “Australian Ordovician Brachiopod
Studies’. Memoirs of the Association of Australasian
Palaeontologists 11, 107-177.
Percival, I.G. (1992). Ordovician brachiopod
biostratigraphy of central-western New South
Wales. In B.D. Webby and J.R. Laurie (eds) “Global
Perspectives on Ordovician Geology (Proceedings of
the 6th International Symposium on the Ordovician
System)’, pp. 215-229. (Balkema, Rotterdam).
Percival, I.G. and Glen, R.A. (2007). Ordovician to
earliest Silurian history of the Macquarie Arc,
Lachlan Orogen, New South Wales. Australian
Journal of Earth Sciences 54, 143-165.
Percival, I.G. and Webby, B.D. (1996). Island benthic
assemblages: with examples from the Late
Ordovician of Eastern Australia. Historical Biology
11, 171-185.
Percival, I.G., Webby, B.D. and Pickett, J.W. (2001).
Ordovician (Bendigonian, Darriwilian to Gisbornian)
faunas from the northern Molong Volcanic Belt of
central New South Wales. Alcheringa 25, 211-250.
Proc. Linn. Soc. N.S.W., 130, 2009
Pickett, J.W. and Percival, I.G. (2001). Ordovician faunas
and biostratigraphy in the Gunningbland area, central
New South Wales. Alcheringa 25, 9-52.
Popov, L.E. and Cocks, L.R.M. (2006). Late Ordovician
brachiopods from the Dulankara Formation of
the Chu-Ili Range, Kazakhstan: their systematics,
palaeoecology and palaeobiogeography.
Palaeontology 49, 247-283.
Popov, L.E., Cocks, L.R.M. and Nikitin, I.F. (2002).
Upper Ordovician brachiopods from the Anderken
Formation, Kazakhstan: their ecology and
systematics. Bulletin Natural History Museum
London (Geology) 58, 13-79.
Popov, L.E., Nikitin, I.F. and Cocks, L.R.M. (2000). Late
Ordovician brachiopods from the Otar Member of the
Chu-Ili Range, south Kazakhstan. Palaeontology 43,
833-870.
Rong, J.-y. (1984). Brachiopods of latest Ordovician
in the Yichang district, western Hubei, central
China. Stratigraphy and Palaeontology of Systemic
Boundaries in China: Ordovician-Silurian boundary
1, 111-176.
Rong, J.-y. and Cocks, L.R.M. (1994). True Strophomena
and a revision of the classification and evolution of
strophomenoid and ‘strophodontoid’ brachiopods.
Palaeontology 37, 651-694.
R6omusoks, A. (1993). Four new brachiopod genera of
the subfamily Oepikinae (Strophomenacea) from the
Ordovician of Estonia. Proceedings of the Academy
of Sciences of the Estonian SSR, Geology 42, 48-57.
R6dmusoks, A. (2004). Ordovician strophomenoid
brachiopods of northern Estonia. Fossilia Baltica 3,
151 pp. + 34 pl.
Rukavishnikova, T.B. (1956). Brakhiopody ordovika Chu-
Tliyskikh gor. [Ordovician brachiopods of the Chu-Ili
Range]. Trudy Geologicheskogo Instituta Akademii
Nauk SSSR 1, 105-168.
Sapelnikov, V.P. and Rukavishnikova, T.B. (1975).
“Verkhneordovikskie, Siluriiskie i Nizhnedevonskie
Pentameridy Kazakhstana’ [Upper Ordovician,
Silurian and Lower Devonian pentamerids of
Kazakhstan]. Akademia Nauk, Ural’skii Nauchnyi
Tsentr, Institut Geologii i Geofiziki, 227 pp. + 43 pls.
(Izdatel’stvo “Nauka”, Sverdlovsk).
Schuchert, C. and LeVene, C.M. (1929). New names for
brachiopod homonyms. American Journal of Science
17, 119-122.
Semeniuk, V. (1970). The Lower-Middle Palaeozoic
stratigraphy of the Bowan Park area, central-western
New South Wales. Journal and Proceedings of the
Royal Society of New South Wales 103, 15-30.
Semeniuk, V. (1973). The stratigraphy of the Bowan Park
Group, New South Wales. Journal and Proceedings
of the Royal Society of New South Wales 105, 77-85.
Temple, J.T. (1965). Upper Ordovician brachiopods from
Poland and Britain. Acta Palaeontologica Polonica
10, 379-427.
Ulrich, E.O. and Cooper, G.A. (1936). New genera and
species of Ozarkian and Canadian brachiopods.
Journal of Paleontology 10, 616-631.
BRACHIOPODS FROM CENTRAL NEW SOUTH WALES
Webby, B.D. (1973). Remopleurides and other Upper
Ordovician trilobites from New South Wales.
Palaeontology 16, 445-475.
Webby, B.D. (1974). Upper Ordovician trilobites from
central New South Wales. Palaeontology 17, 203-
Dar,
Webby, B.D. and Morris, D.G. (1976). New Ordovician
stromatoporoids from New South Wales. Journal and
Proceedings of the Royal Society of New South Wales
109, 125-135.
Webby, B.D. and Packham, G.H. (1982). Stratigraphy and
regional setting of the Cliefden Caves Limestone
Group (Late Ordovician), central-western New South
Wales. Journal of the Geological Society of Australia
29, 297-317.
Williams, A., Brunton, C.H.C. and Carlson, S.J. (eds).
(2000). ‘Treatise on Invertebrate Paleontology, Part
H. Brachiopoda (Revised), Vol. 3’. (The Geological
Society of America: Boulder, and The University of
Kansas Press: Lawrence).
Zhan, R.-b., Jin, J., Rong, J.-y., Chen, P.-f. and Yu, G.-
h. (2008). Strophomenide brachiopods from the
Changwu Formation (Late Katian, Late Ordovician)
of Chun’an, western Zheijiang, south-east China.
Palaeontology 51, 737-766.
Zhen, Y.-y. and Pickett, J.W. (2008). Ordovician (Early
Darriwilian) conodonts and sponges from west of
Parkes, central New South Wales. Proceedings of the
Linnean Society of New South Wales 129, 57-82.
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
* BILLABONG
CREEK
~ LIMESTONE
e e PARKES
Gunningbland
e FORBES
® Evugowra
NEW SOUTH WALES
Bathurst
| . SYDNEY
<
400km
N
Molong |
SEEDY CREEK
i LIMESTONE
U
Cudal
e ORANGE e
Ww
— sowan PARK
CARGO CREEK pamRP ci
LIMESTONE ,
\ Cargo
: _— REGANS CREEK
~ LIMESTONE
! ¢
CANOMODINE — \ -
LIMESTONE
® Canowindra
Sia
Licking Hole Creek —
CLIEFDEN CAVES
+— LIMESTONE
~~ SUBGROUP
6
Mandurama
® \Voodstock
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.
REFERENCES
Balashov, Z.G. (1955). Klass Cephalopoda. Otryad
Nautiloidea [Class Cephalopoda. Order Nautiloidea].
In Nikiforova, O.I. (ed.) Polevoi Atlas, Ordovikski
i siluriiski fauny Sibirskoi Platformy, 87-104.
(VSEGEI: Moscow).
Balashov, Z.G. (1962). Nautiloidei Ordovika Sibirskoi
Platformy [Ordovician Nautiloids of the Siberian
Platform]. /zdatelstvo, Leningradskogo Universiteta,
205 pp.
Billings, E. (1859). Nautilus jason. Canadian Naturalist
and Geologist 4, 464.
Billings, E. (1865). ‘Palaeozoic Fossils: containing
description and figures of new and little known
species of organic remains from Silurian rocks.
189
RARE FOSSILS FROM THE LATE ORDOVICIAN
Volume 1’. Geological Survey of Canada. 426 pp.
(Dawson Brothers, Montreal).
Bischoff, G.C.O. (1973). On the nature of the conodont
animal. Geologica et Palaeontologica 7, 147-174.
Bischoff, G.C.O. (1978). Internal structures of conulariid
tests and their functional significance, with special
reference to Circonulariina n. suborder (Cnidaria,
Scyphozoa). Senckenbergiana lethaea 59, 275-327.
Cope, J.C.W. (2004). Bivalve and rostroconch mollusks.
In ‘The Great Ordovician Biodiversification Event’
(Eds B.D. Webby, F. Paris, M.L. Droser and I.G.
Percival) pp. 196-208. (Columbia University Press,
New York).
Edgecombe, G.D. and Webby, B.D. (2006). The
Ordovician encrinurid trilobite Sinocybele from New
South Wales and its biogeographic significance.
Memoirs of the Association of Australasian
Palaeontologists 32, 413-422.
Edgecombe, G.D. and Webby, B.D. (2007). Ordovician
trilobites with eastern Gondwanan affinities from
central-west New South Wales and Tasmania.
Memoirs of the Association of Australasian
Palaeontologists 34, 255-281.
Foerste, A.F. (1928). American Arctic and related
cephalopods. Denison University Bulletin, Journal of
the Scientific Laboratories 23, 1-110, 29 pls.
Frey, R.C. (1995). Middle and Upper Ordovician Nautiloid
Cephalopods of the Cincinnati Arch region of
Kentucky, Indiana and Ohio. U.S. Geological Survey
Professional Paper 1066-P, 1-126, 22 pls.
Furnish, W.M. and Glenister, B.F. (1964). Nautiloidea—
Tarphycerida. In “Treatise on Invertebrate
Paleontology, Part K. Mollusca 3’ (Eds C. Teichert,
et al.) pp. K343-K368. (Geological Society of
America: New York, and University of Kansas Press:
Lawrence).
Glenister, B.F. (1952). Ordovician nautiloids from New
South Wales. Australian Journal of Science 15, 89-
91.
Jerre, F. (1993). Conulariid microfossils from the
Silurian Lower Visby Beds of Gotland, Sweden.
Palaeontology 36, 403-424.
Jerre, F. (1994). Anatomy and phylogenetic significance of
Eoconularia loculata, a conulariid from the Silurian
of Gotland. Lethaia 27, 97-109.
Ladd, H.S. (1929). The stratigraphy and paleontology
of the Maquoketa Shale of Iowa. Jowa Geological
Survey Annual Report 34, 307-448. (not seen — fide
Van Iten et al. 1996).
Laurie, J.R. (1991). Articulate brachiopods from the
Ordovician and Lower Silurian of Tasmania. In
P.A. Jell (ed.) Australian Ordovician Brachiopod
Studies. Memoirs of the Association of Australasian
Palaeontologists 11, 1-106.
Leme, J.M., Heredia, S., Rodrigues, S.C., SimGes,
M.G., Acefiolaza, G.F. and Milana, J.P. (2003).
Teresconularia gen. nov. from the Lower Ordovician
of the Cordillera Oriental of Salta (NW Argentina):
the oldest conulariid (Cnidaria) from South America.
190
Revista Espanola de Micropaleontologia 35, 265-
DP.
Leme, J.M., Simdes, M.G., Marques, A.C. and Van
Iten, H. (2008). Cladistic analysis of the suborder
Conulariina Miller and Gurley, 1896 (Cnidaria,
Scyphozoa; Vendian — Triassic). Palaeontology 51,
649-662.
Lindstrom, G. (1884). On the Silurian Gastropoda
and Pteropoda of Gotland. Svenska Vetenskaps-
Akademiens Handlingar 19, 1-250. (not seen — fide
Jerre 1993).
Parfrey, S.M. (1982). Palaeozoic conulariids from
Tasmania. Alcheringa 6, 69-75.
Percival, I.G. (1976). The geology of the Licking Hole
Creek area, near Walli, central western New South
Wales. Journal and Proceedings of the Royal Society
of New South Wales 109, 7-23.
Percival, I.G. (1978). Inarticulate brachiopods from the
Late Ordovician of New South Wales, and their
palaeoecological significance. Alcheringa 2, 117-141.
Percival, I.G. (1979a). Ordovician plectambonitacean
brachiopods from New South Wales. Alcheringa 3,
91-116.
Percival, I.G. (1979b). Late Ordovician articulate
brachiopods from Gunningbland, central western
New South Wales. Proceedings of the Linnean
Society of New South Wales 103, 175-187.
Percival, I.G. (2005). Late Ordovician deepwater (BA 4)
brachiopods of circum-Pacific terranes. In Abstracts
for the Second International Symposium of IGCP503
on Ordovician Palaeogeography and Palaeoclimate,
June 15-18, 2005. Insight (a Milwaukee Public
Museum series in Natural History) 2, 24-25.
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.
Percival, I.G., Engelbretsen, M.J. and Brock, G.A. (1999).
Distribution of Middle to Late Ordovician lingulate
brachiopods in New South Wales. Acta Universitatis
Carolinae, Geologica 43, 347-350.
Percival, I.G. and Webby, B.D. (1996). Island benthic
assemblages: with examples from the Late
Ordovician of Eastern Australia. Historical Biology
11, 171-185.
Percival, I.G., Zhen Y.-Y. and Pickett, J.W. (2006). Late
Ordovician faunas from the Quandialla-Marsden
district, south-central New South Wales. Proceedings
of the Linnean Society of New South Wales 127, 235-
DIS).
Pickett, J.W. and Percival, I.G. (2001). Ordovician faunas
and biostratigraphy in the Gunningbland area, central
New South Wales. Alcheringa 25, 9-52.
Pojeta, J. jr., Gilbert-Tomlinson, J. and Shergold, J.H.
(1977). Cambrian and Ordovician rostroconch
mollusks from Northern Australia. Bureau of Mineral
Resources, Geology and Geophysics, Bulletin 171, 54
pp. + 27 pls.
Proc. Linn. Soc. N.S.W., 130, 2009
I.G. PERCIVAL
Pojeta, J. jr. and Runnegar, B. (1976). The paleontology Zhen Y.-Y., Webby B.D. and Barnes C.R. (1999). Upper
of rostroconch mollusks and the early history of the Ordovician conodonts from the Bowan Park
Phylum Mollusca. United States Geological Survey succession, central New South Wales, Australia.
Professional Paper 968, 88 pp. + 54 pls. Géobios 32, 73-104.
Popov, L.E., Cope, J.C.W. and Nikitin, I.F. (2003). A new
Ordovician rostroconch mollusc from Kazakhstan.
Alcheringa 27, 173-179.
Rong J.-y., Zhan R.-b. and Han N.-r. (1994). The oldest
known Eospirifer (Brachiopoda) in the Changwu
Formation (Late Ordovician) of western Zhejiang,
east China, with a review of the earliest spiriferoids.
Journal of Paleontology 68, 763-776.
Semeniuk, V. (1973). The stratigraphy of the Bowan Park
Group, New South Wales. Journal and Proceedings
of the Royal Society of New South Wales 105, 77-85.
Sherwin, L. (1970). Silurian conularids from New South
Wales. Records of the Geological Survey of New
South Wales 11, 35-41.
Sowerby, J. (1821). ‘The mineral conchology of Great
Britam; or coloured figures and descriptions of those
remains of testaceous animals or shells, which have
been preserved at various times, and depths in the
earth’. 194 pp. (W. Arding Co., London).
Stait, B.A., Webby, B.D. and Percival, I.G. (1985). Late
Ordovician nautiloids from central New South Wales,
Australia. Alcheringa 9, 143-157.
Van Iten, H., Fitzke, J.A. and Cox, R.S. (1996).
Problematical fossil cnidarians from the Upper
Ordovician of the north-central USA. Palaeontology
39, 1037-1064.
Van Iten, H. and Vyhlasova, Z. (2004). Conulartids. In
“The Great Ordovician Biodiversification Event’ (Eds
B.D. Webby, F. Paris, M.L. Droser and I.G. Percival)
pp. 119-123. (Columbia University Press, New York).
Webby, B.D. (1992). Ordovician island biotas: New South
Wales record and global implications. Journal and
Proceedings of the Royal Society of New South Wales
125, 51-77.
Webby, B.D. and Blom, W.M. (1986). The first well-
preserved radiolarians from the Ordovician of
Australia. Journal of Paleontology 60, 145-157.
Webby, B.D. and Morris, D.G. (1976). New Ordovician
stromatoporoids from New South Wales. Journal and
Proceedings of the Royal Society of New South Wales
109, 125-135.
Webby, B.D. and Packham, G.H. (1982). Stratigraphy and
regional setting of the Cliefden Caves Limestone
Group (Late Ordovician), central-western New South
Wales. Journal of the Geological Society of Australia
29, 297-317.
Webby, B.D. and Trotter, J. (1993). Ordovician sponge
spicules from New South Wales, Australia. Journal of
Paleontology 67, 28-41.
Wiman, C. (1895). Palaeontologische Notizen 1—2.
Bulletin of the Geological Institution of Upsala 3(2),
1-9. (not seen — fide Jerre 1994).
Proc. Linn. Soc. N.S.W., 130, 2009 191
veocget st ROL err I ‘al tea
{thee pean otha seeggnad suntan w
», eed reas Ay pa, ae eee
yi ~Y he
in aeari ne ah i a
nd thei The "nt m
rence 4 tae (PLB PE Nar
Seypharow). Seni Sugeber ged el alee any Sep
WA fuk Un ‘ erie a OS 4a
ih iy Leet ian Pi Aa
ti , F fn wad ek
Ny ¥ r
1
“f "ot
i ; *
i”
rv
¥
ay i. we
wey AH) hs
ie as al
“We ii
ee
'
ra . Lod U
vt 4 " ot
At pr ry
1 die Wi ™~ 4
Hier, Bald i
Sounds Wieh 4
41
i B, CC
bat ati * Vins
erticy , “4
yt ve wticitry pal
te ser oa
i ee 1
: BAT a arp tay ae hg
¥ oh'va Sh ‘, it, ny i
Ti Wl ee ia it 4:08 at |
i a je
am 1 ECW ate oe gbargaiat Hepp ce:
r ane? Loker’ So et nv Tienes Hi
i ted, on DP tay Ceri firey : nepal
‘4 fetralrs ay ihe Aveoehelon Of Abarnimem
‘ in * hI Pe eMe ;
* ; ei S.. ochenpiben Cy Breed
1 Ki cartels | ET) aie rad, dP Ct f
pil eire ally abl tit Pivevat fie |, SHOT i wdo vinings
oH tic Com tAieog alia NW Arypesctiow): .
trad yh, xvarierial wrist niet fromm, Sabet | Naericn 0,
if 9
stein “ihe BAA pes an td ae
» Mens Mihara, a wal’ Boos
vedio
Oey roy NY
wversstrond den sun ts taaiapreiabenntl
" yetaphigts sro dh Anoka :
rio itine Heitey stT To Mere, thi sha it
wey? aa ety: ER He segeabarnanaleeh
set patel Ue y depamgsabtirde aah aren ye a
vefipetine then") bush: oq ok-aalgea Aiba 193
Ct Feat ated wnt 1 st iN
Ase, ah iestehneunos tap oo é
wi Ne aero araanaiey?. a 6 aaah, 2
val: £80. APR TRY, Yeonwetteal ode heeiee ae wy
tenn 1 iccavhaibevsiaxoahin gas aid ia.
er art we ap a
pray ep vita
ytd ete pts 2 fatica MI, an bat
a fe} 1 dda da lh sei
el ria sealfelunl aaamaa shan} ee al
Se Soatiy Wake PAL Rrmapeioarlalty,
avy (RPO. KPa kero ah | peal
rae Coie sie v4
“Yor He Semon Porcine
eh Pieters
aba} ane tao RE 7
(dn Waa yok a 290°0N Ride a
aay wie a emai icra a BLL
hace wort jam ively
RTT heat yah, sautaghepuiciotatn Wil
eh Hi, haan yop Loonie ago yy}
es
=
tow ion ot AOROL nA, Wo redthk ;
vali ‘iconanedon ant omelet
CGAP) eb etgehayneycus) oa a cl
sahuneesgcel niblaie 0 te. fi aiephhiliad
ity yeaa ins. taal eope iy
o ehh ahha wih UE manors |
stance
3 Uiditin aera!
ites Mast yey pnb in’
Otway moan, ,
VO Vek ee
ano reign, ie sie eh in iG
“or bana meet nny
fot 4 sai
Sivne
A og
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
New England] med Karikeree Metadolerite
Fald Belt ~—4 ANY 8 ee iene
A Thrumster Slate Roads
— —— =Faits
[| water ee
N
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
GDA 1994 - MGA Zone 56
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.
REFERENCES
Breyvel’, M.G., Bogoyavlensaya, O.V., Breyvel’, I.A.,
Khodalevich, A.N., Shurygina, M.V., & Yanet, F.E.M.
(1972). Kishepolostnye 1 brakhiopody zhivetskikh
otlozheniy vostochnogo sklona Urala. Nedra,
Moscow, 264 p.
Brown, I.A. (1942). The Tamworth Series (Lower and
Middle Devonian) near Attunga, N.S.W. Journal and
Proceedings of the Royal Society of New South Wales
75, 165-176.
Briihl, D., & Pohler, S. (1999). Tabulate corals from
the Moore Creek Limestone (Middle Devonian:
Late Eifelian-Early Givetian) in the Tamworth Belt
(New South Wales, Australia). Abhandlungen der
Geologischen Bundesanstalt 54, 275-293.
Bultynck, P. (1987). Pelagic and neritic conodont
successions from the Givetian of pre-Sahara Morocco
and the Ardennes. Bu/letin de I Institut royal des
Sciences naturelles de Belgique, Sciences de la Terre
57, 149-181.
Campbell, K.S.W. (1957). A Lower Carboniferous
brachiopod — coral fauna from New South Wales.
Journal of Paleontology 31(1), 34-98, pl. 11-17.
Cawood, P. A. (1983). Modal composition and
clinopyroxene geochemistry of lithic sandstones
from the New England Fold Belt (east Australia):
a Paleozoic fore-are terrain. Geological Society of
America Bulletin 94, 1199-1214.
Cawood, P. A., & Leitch, E. C. (1985). Accretion and
dispersal tectonics of the southern New England
Fold Belt, eastern Australia, in Howell, D. G., ed.
“Tectonostratigraphic terranes of the circum-Pacific
region’, Circum-Pacific Council for Energy and
Mineral Resources Earth Science Series, Number 1,
481-492, Houston, Texas.
Chapman, F. (1907). New Silurian fossils of eastern
Victoria, Part 1. Records of the Geological Survey of
Victoria 2, 67-80.
Chapman, F. (1914). Newer Silurian fossils of eastern
Victoria - Part III. Records of the Geological Survey
of Victoria 3, 301-316, pl. 46-61.
Chapman, F. (1918). Appendix 2. Note on a new species
ZW
of Chaetetes. In Benson, W.N., The geology and
petrology of the Great Serpentine Belt of New South
Wales. Part VII. The geology of the Loomberah
district and a portion of the Goonoo Goonoo estate.
Proceedings of the Linnean Society of New South
Wales 43, 392-394, pl. 42.
Chapman, F. (1920). Appendix. Lower Carboniferous
limestone fossils from New South Wales. In Benson,
W.N., Dun, W.S., & Browne, W.R., The geology
and petrology of the Great Serpentine Belt of New
South Wales. Part IX. The geology, palaeontology
and petrography of the Currabubula district, with
notes on adjacent regions. Section B. Palaeontology.
Proceedings of the Linnean Society of New South
Wales 45, 364-367, pl. 24.
Chapman, F. (1921). New or little-known Victorian fossils
in the National Museum. Part XXV. Some Silurian
tabulate corals. Proceedings of the Royal Society of
Victoria 33, 212-225, pl. 9-11.
Chapman, F. (1925). New or little-known fossils in the
National Museum. Part XX VIII. Some Silurian
rugose corals. Proceedings of the Royal Society of
Victoria 37, 104-118, pl. 12-15.
Chernyshev, B.B. (1937). Siluriyskie i devonskie Tabulata
Mongolii 1 Tuvy. Trudy Akademii nauka SSSR,
Mongol skogo Komiteta, 30(6), 1-31, pl. 1-4.
Chernyshev, B.B. (1941). Siluriyskie i nizhnedevonskie
korally basseyna reki Tarei (yugo-zapadnoy Taymyr).
Trudy vsesoyuznogo Arkticheskogo Instituta 158(5),
9-64.
Clausen, C.-D., Leuteritz, K., & Ziegler, W. (1979).
Biostratigraphie und Lithofazies am Siidrand
der Elsper Mulde (hohes Mittel- und tiefstes
Oberdevon; Sauerland, Rheinisches Schiefergebirge).
Geologisches Jahrbuch AS51, 3-37, | pl.
Dana, J.D. (1846). Structure and classification of
zoophytes. “U.S. Exploring Expedition during the
years 1838 — 1842 under the command of Charles
Wilkes, U.S.A.’ 7, x + 740 p. Lea & Blanchard,
Philadelphia.
de Blainville, H.M.D. (1830). Zoophytes. In Dictionnaire
des Sceinces naturelles 60. 1-546 (not seen).
D’Orbigny, A. (1849). “Note sur des polypiers fossiles.’
1-12. Victor Masson, Paris.
Dubatoloy, V.N. (1963). “Pozdnesiluriyskie i devonskie
tabulyaty, geliolitidy 1 khetetidy Kuznetzkogo
basseyna.’ 193 p. Nauka, Moscow.
Duncan, P.M. (1872). Third report on the British fossil
corals. Report 41 Meeting British Association for the
Advancement of Science, Edinburgh (1871), 116-137.
Dybowski, W.N. (1873). Monographie der Zoantharia
Sclerodermata Rugosa aus der Silurformation
Estlands, Nord-Livlands und der Insel Gotland.
Archiv fiir Naturkunde Liv-, Ehst- und Kurlands, ser:
1,5, 257-414, pl. 1-2.
Eichwald, C.E. von (1829). “Zoologia specialis quam
expositis animalibus tum vivis, tum fossilibus
potissimum Rossiae in universum, et Poloniae in
specie, in usum lectionum.’ Vol. 1, vi+ 314 p., 5 pl. J.
Zawalski, Vilna.
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
Etheridge, R., Jr (1890). Descriptions of Upper Silurian
fossils from the Lilydale limestone, Upper Yarra
district, Victoria. Records of the Australian Museum
1, 60-67, pl. 8-9.
Etheridge, R., Jr (1892). Descriptions of four
Madreporaria Rugosa - species of the genera
Phillipsastrea, Heliophyllum and Cyathophyllum
- from the Palaeozoic rocks of New South Wales.
Records of the Geological Survey of New South Wales
2, 165-174, pl. 11-12.
Etheridge, R., Jr (1896). Description of a small collection
of Tasmanian Silurian fossils presented to the
Australian Museum by Mr A. Montgomery, M.A.,
Government Geologist, Tasmania. Report of the
Secretary of Mines Tasmania, xli-xlviii, | pl.
Reprinted 1897, Papers and Proceedings of the Royal
Society of Tasmania 1896, 29-46, pl. 1.
Etheridge, R., Jr (1899). On the corals of the Tamworth
district, chiefly from the Moore Creek and Woolomol
Limestones. Records of the Geological Survey of New
South Wales 6, 151-182, pl. 16-38.
Etheridge, R., Jr (1902). Additions to the Middle Devonian
and Carboniferous corals in the Australian Museum.
Records of the Australian Museum 4, 253-262, pl.
37-40.
Etheridge, R., Jr (1911). The Lower Palaeozoic corals of
Chillagoe and Clermont. Part 1. Publications of the
Geological Survey of Queensland 231, 1-8, pl. A-D.
Etheridge, R., Jr (1917). Descriptions of some Queensland
Palaeozoic and Mesozoic fossils. 4, Vetofistula, a new
form of Palaeozoic Polyzoa, allied to Rhabdomeson
Young & Young, from Reid’s Gap, near Townsville.
Publications of the Geological Survey of Queensland
260, 17-29, pl. 4.
Etheridge, R., Jr (1921). Palaeontologia Novae Cambriae
Meridionalis — Occasional descriptions of New South
Wales fossils. No. 8. Records of the Geological
Survey of New South Wales 10, 1-11, pl. 1-7.
Etheridge, R., Jr & Foord, A.H. (1884). On two species
of Alveolites and one of Amplexopora from the
Devonian rocks of northern Queensland. Annals and
Magazine of Natural History (5) 14, 175-179, pl. 6.
Finks, R.M., Reid, R.E.H., & Rigby, J.K. (2004). “Treatise
on Invertebrate Paleontology, ed. C. Teichert. Part E,
Porifera (Revised).’ Volume 3. Geological Society of
America and University of Kansas.
Fitzgerald, J.K. (1955). A new tabulate coral from New
South Wales. Journal of Paleontology 29, 1057-1059,
1 text-fig.
Gerth, H. (1921). Die Anthozoen der Dyas von Timor.
Paldontologie von Timor 9(16), 65-147, pl. 145-150.
Goldfuss, G.A. (1826). ‘Petrefacta Germaniae.’ 1, 1-76.
Armz & Co., Diisseldorf.
Hall, J. (1851). New genera of fossil corals from the report
by James Hall, on the palaeontology of New York.
American Journal of Science, series 2, 11, 398-401.
Hartman, W.D., & Goreau, T.F. (1972). Ceratoporella
(Porifera: Sclerospongiae) and the chaetetid “corals”.
Transactions of the Connecticut Academy of Arts and
Sciences 44, 133-148.
Proc. Linn. Soc. N.S.W., 130, 2009
Hill, D. (1939). The Middle Devonian rugose corals of
Queensland, I. Douglas Creek and Drummond Creek,
Clermont district. Proceedings of the Royal Society of
Queensland 50, 55-65, pl. 4-5.
Hill, D. (1940a). The Middle Devonian rugose corals of
Queensland, II. The Silverwood — Lucky Valley area.
Proceedings of the Royal Society of Queensland 51,
150-168, pl. 2-3.
Hill, D. (1940b). The Lower Devonian rugose corals of
the Murrumbidgee and Goodradigbee Rivers, N.S.W.
Journal and Proceedings of the Royal Society of New
South Wales 74, 247-276, pl. 9-11.
Hill, D. (1942a). The Middle Devonian rugose corals of
Queensland, III. Burdekin Downs, Fanning R., and
Reid Gap, North Queensland. Proceedings of the
Royal Society of Queensland 53, 229-268, pl. 5-11.
Hill, D. (1942b). The Lower Devonian rugose corals from
the Mt. Etna Limestone, Qld. Proceedings of the
Royal Society of Queensland 54, 13-22, pl. 1.
Hill, D. (1942c). The Devonian rugose corals of the
Tamworth district, N.S.W. Journal and Proceedings
of the Royal Society of New South Wales 76, 142-164,
pl. 2-4.
Hill, D. (1942d). Middle Palaeozoic rugose corals from the
Wellington district, N.S.W. Journal and Proceedings
of the Royal Society of New South Wales 76, 182-189,
pl. 5-6.
Hill, D. (1950). Middle Devonian corals from the Buchan
district, Victoria. Proceedings of the Royal Society of
Victoria 62, 137-164, pl. 5-9.
Hill, D. (1954). Devonian corals from Waratah Bay,
Victoria. Proceedings of the Royal Society of Victoria
66, 7-118, pl. 6-9.
Hill, D. (1955). Ordovician corals from Ida Bay,
Queenstown and Zeehan, Tasmania. Papers and
Proceedings of the Royal Society of Tasmania 89,
237-254, pl. 1-3.
Hill, D. (1957). Ordovician corals from New South Wales.
Journal and Proceedings of the Royal Society of New
South Wales 91, 97-107, pl. 2-4.
Hill, D. (1981). ‘ Treatise on Invertebrate Paleontology,
ed. C. Teichert. Part F, Coelenterata, Supplement 1,
Rugosa and Tabulata.’ 2 vols. Geological Society of
America and University of Kansas.
Hill, D., & Jones, O.A. (1940). The corals of the Garra
Beds, Molong district, New South Wales. Journal
and Proceedings of the Royal Society of New South
Wales 74, 175-208, pl. 2-8.
Hill, D., Playford, G., & Woods, J.T. (1967). °
Devonian fossils of Queensland.’ Queensland
Palaeontographical Society, Brisbane. 1-32, pl. 1-15.
Hinde, G.J. (1890). Notes on the palaeontology of
Western Australia, 2. Corals and Polyzoa. Geological
Magazine (3) 7, 194-204, pl. 8, 8a.
Hooper, J.N.A., & Van Soest, R.W.M., eds (2002).
‘Systema Porifera: a guide to the classification
of sponges.’ 2 vols. Kluwer Academic/Plenum
Publishers, New York.
Ishiga, H., Leitch, E.C., Watanabe, T., Naka, T., &
Iwasaki, M. (1988). Radiolarian and conodont
23
DEVONIAN MARINE INVERTEBRATE FOSSILS
biostratigraphy of siliceous rocks from the New
England Fold Belt. Australian Journal of Earth
Sciences 35(1), 73-80.
Jell, J.S., & Hill, D. (1969). Devonian corals from the
Ukalunda district, north Queensland. Publications
of the Geological Survey of Queensland 340
(Palaeontological Papers No. 16), 1-27, pl. 1-9.
Jell, J.S., & Hill, D. (1970a). The Devonian coral fauna
of the Point Hibbs Limestone, Tasmania. Papers and
Proceedings of the Royal Society of Tasmania 104,
1-16, pl. 1-6.
Jell, J.S., & Hill, D. (1970b). Revision of the coral fauna
from the Devonian Douglas Creek Limestone,
Clermont, central Queensland. Proceedings of the
Royal Society of Queensland 81(10), 93-120, pl. 3-8.
Jones, O.A. (1937). The Australian massive species of
the coral genus Favosites. Records of the Australian
Museum 20, 79-102, 11-16.
Jones, O.A. (1941). The Devonian Tabulata of Douglas
and Drummond Creeks, Clermont, Queensland.
Proceedings of the Royal Society of Queensland
53(3), 41-60, pl. 1-3.
Koninck, L.G. de (1876-77). “Recherches sur les
fossils paléozoiques de la Nouvelle-Galles du Sud
(Australie).’ 1-373, Atlas (pl. 1-4, 1876; pl. 5-25,
1877) Brussels. [Re-issued in 1877-1878 in Mémoires
de la Société royale des Sciences de Liége (2) 6(2),
1-140, pl. 1-4 (1877); 7, 1-235, pl. 5-24 (1878). An
English translation, Descriptions of the Palaeozoic
fossils of New South Wales, differently paged but
with the same plates, was issued in 1898 as Memoirs
of the Geological Survey of New South Wales,
Palaeontology 6, \-xiii + 1-298, pl. 1-24.
Lamarck, J.B.P.A. de M. (1801). “Systeme des animaux
sans vertebres.’ villi + 432 p. The author, Paris.
Lamarck, J.B.P.A de M. (1816). ‘Histoire naturelle des
animaux sans vertebres.’ 2, 1-568. The author, Paris.
Lecompte, M. (1933). Le genre A/veolites Lamarck
dans le Dévonien moyen et supérieur de |’ Ardenne.
Mémoires du Muséum royale d'Histoire naturelle de
Belgique 55, 1-50, pl. 1-4.
Lecompte, M. (1939). Les tabulés du Dévonien moyen et
supérieur du bord sud du Bassin de Dinant. Mémoires
du Muséum royale d'Histoire naturelle de Belgique
90, 1-229, pl. 1-23.
Leitch, E.C. (1980). Rock units, structure and
metamorphism of the Port Macquarie Block, eastern
New England Fold Belt. Proceedings of the Linnean
Society of New South Wales 104(4), 273-292.
Lindstrém, G. (1876). On the affinities of the Anthozoa
Tabulata. Annals and Magazine of Natural History,
ser. 4, 18, 1-17.
Lonsdale, W. (1840). In Sedgwick, A., & Murchison, R.L.,
On the physical structure of Devonshire, and on the
subdivisions and geological relations of its older
stratified deposits. Transactions of the Geological
Society of London, ser. 2, 5, p.697.
Mawson, R., & Talent, J.A. (1989). Late Emsian
— Givetian stratigraphy and conodont biofacies
— carbonate slope and offshore shoal to sheltered
214
lagoon and nearshore carbonate ramp — Broken River,
North Queensland, Australia. Courier Forschungs-
Institut Senckenberg 117, 205-259.
Mawson, R., & Talent, J.-A. (1994a). The Tamworth
Group (mid-Devonian) at Attunga, New South
Wales: conodont data and inferred ages. Courier
Forschungs-Institut Senckenberg 168, 37-59.
Mawson, R., & Talent, J.A. (1994b). Age of an Early
Devonian carbonate fan and isolated limestone clasts
and megaclasts, east-central Victoria. Proceedings of
the Royal Society of Victoria 106, 31-70.
Mawson, R., & Talent, J.A. (2000). The Devonian of
eastern Australia: stratigraphic alignments, stage and
series boundaries, and the transgresssion-regression
pattern re-considered. Courier Forschungs-Institut
Senckenberg 225, 243-270.
Mawson, R., & Talent, J.A. (2003). Conodont faunas
from sequences on or marginal to the Anakie Inlier
(Central Queensland, Australia) in relation to
Devonian transgressions. Bulletin of Geosciences
78(4), 335-358.
Milne-Edwards, H., & Haime, J. (1850). A monograph of
the British fossil corals, p. 1— Ixxxv, 1-71, pl. 1-11.
Palaeontographical Society Monographs, London.
Milne-Edwards, H., & Haime, J. (1851). Monographie des
polypiers fossils des terrains paléozoiques. Archives
du Muséum Nationale d'Histoire naturelle 5, 1-502,
pl. 1-20.
Moore, D., & Roberts, J. (1976). The Early Carboniferous
marine transgression in the Merlewood Formation,
Werrie Syncline, New South Wales. Journal and
Proceedings of the Royal Society of New South Wales
109(1-2), 49-57.
Nicholson, H.A., & Etheridge, R., Jr (1879). Descriptions
of Palaeozoic corals from northern Queensland, with
observations on the genus Stenopora. Annals and
Magazine of Natural History (5), 4, 216-226, 265-
285, pl. 14.
Och, D.J., Percival, I.G., & Leitch, E.C. (2007).
Ordovician conodonts from the Watonga Formation,
Port Macquarie, northeast New South Wales.
Proceedings of the Linnean Society of New South
Wales 128, 209-216.
Oliver, W.A. (1963). Redescription of three species of
corals from the Rockport Dolomite of New York.
U.S.G.S. Professional Paper 414G, 1-9, pl. 1-5.
Pedder, A.E.H. (1964). Two new genera of Devonian
tetracorals from Australia. Proceedings of the
Linnean Society of New South Wales 88, 364-367, pl.
19.
Pedder, A.E.H. (1965). A revision of the Australian
Devonian corals previously referred to Mictophyllum.
Proceedings of the Royal Society of Victoria 78(2),
201-220, pl, 30-34.
Pedder, A.E.H. (1966). The Devonian tetracoral
Haplothecia and new Australian phacellophyllids.
Proceedings of the Linnean Society of New South
Wales 90, 181-189, pl. 6.
Pedder, A.E.H., Jackson, J.H., & Philip, G.M. (1970a).
Lower Devonian biostratigraphy in the Wee Jasper
Proc. Linn. Soc. N.S.W., 130, 2009
J. PICKETT, D. OCH AND E. LEITCH
region of New South Wales. Journal of Paleontology
44, 206-251, pl. 37-50.
Pedder, A.E.H., Jackson, J.H., & Ellenor, D.W. (1970b).
An interim account of the Middle Devonian Timor
Limestone of north-eastern New South Wales.
Proceedings of the Linnean Society of New South
Wales 94(3), 242-272, pl. 14-24.
Philip, G.M. (1960). The Middle Palaeozoic squamulate
favositids of Victoria. Palaeontology 3, 186-207, pl.
30-40.
Philip. G.M. (1962). The palaeontology and stratigraphy
of the Siluro-Devonian sediments of the Tyers area,
Gippsland, Victoria. Proceedings of the Royal Society
of Victoria 75, 123-246, pl. 11-36.
Pickett, J.W. (1967). Lower Carboniferous coral faunas
from the New England District of New South Wales.
Geological Survey of N.S.W., Memoir Palaeontology
15, 1-38, pl. 1-20.
Pickett, J.W. (1969). Middle and Upper Palaeozoic
sponges from New South Wales. Geological Survey
of N.S.W., Memoir Palaeontology 16, 1-25, pl. 1-11.
Pickett, J.W. (1985). A mid-Devonian limestone near
Wauchope. Geological Survey of New South Wales
- Unpublished Palaeontological Report 1985/4 (GS
1985/118).
Pickett, J.W. (1991). Fossil corals from the Touchwood
Formation near Port Macquarie. Geological Survey
of New South Wales - Unpublished Palaeontological
Report 1991/1 (GS 1991/105).
Pickett, J.W. (1999). “OZCORALS: a Bibliography and
index of fossil corals from Antarctica, Australia,
New Guinea and New Zealand.’ Microsoft Access®
database, downloadable from website of Association
of Australasian Palaeontologists.
Pickett, J.W., & Ingpen, I.A. (1990). Ordovician and
Silurian strata south of Trundle, New South Wales.
Quarterly Notes of the Geological Survey of New
South Wales 78, 1-14.
Pickett, J.W., & McClatchie, L. (1991). Age and relations
of stratigraphic units in the Murda Syncline area.
Quarterly Notes of the Geological Survey of New
South Wales 85, 9-32.
Poéta, F. (1902). Anthozoaires et Aleyonaires. In:
Barrande, J., ed., “Systeme silurien du centre de la
Boheme.’ Part 1, 8(2), 1-347. Prague, Paris.
Pohler, S.M.L. (1998). Devonian carbonate buildup facies
in an intra-oceanic island arc (Tamworth Belt, New
South Wales, Australia). Facies 39, 1-34.
Pohler, S.M.L. (2001). Paleoecology, biostratigraphy and
paleogeography of Favositidae (Tabulata) from the
Emsian to Middle Devonian Tamworth Group (New
South Wales, Australia). Senckenbergiana lethaea
81(1), 91-109.
Pohler, S.M.L. (2002). Favositidae (Tabulata)
from Emsian to Middle Devonian limestones
of the Tamworth Group (N.S.W., Australia).
Paldontologische Zeitschrift 76(1), 1-19.
Roberts, J. & Geeve, R. (1999). Allochthonous forearc
blocks and their influence on an orogenic timetable
for the Southern New England Orogen, in Flood,
Proc. Linn. Soc. N.S.W., 130, 2009
P. G., ed. “Regional Geology Tectonics and
Metallogenesis New England Orogen’, 105-114,
University of New England, Armidale.
Roberts, J. , Leitch, E.C. , Lennox, P.G. , & Offler, R.
(1995). Devonian-Carboniferous stratigraphy of the
southern Hastings Block, New England Orogen,
eastern Australia. Australian Journal of Earth
Sciences 42(6), 609-634
Schliiter, C. (1899). Die Anthozoen des rheinischen
Mitteldevons. Abhandlungen zur geologischen
Specialkarte von Preussen und den Thiiringischen
Staaten 8(4), x + 207, pl. 1-16.
Smith, S. (1933). Sur les espéces nouvelles d’ Alveolites
de |’Eifelien inférieur du Nord de la France et de la
Belgique. Annales de la Société géologique du Nord
57, 134-145.
Sokolov, B.S. (1952). Tabulyaty paleozoya evropeyskoy
chasti SSSR, chast’ 4, Devon Russkoy platformy 1
zapadnogo Urala. Trudy Vsesoyuznogo Neftyanogo
Nauchno-issledovannogo geolog-razvedochnogo
Instituta (VNIGRI), n.s. 62, 1-292.
Sokolov, B.S. (1955). Tabulyaty paleozoya evropeyskoy
chasti SSSR. Vvedenie: Obshchie voprozy k
sistematiki i istorii razvitiya tabulyat. Trudy
Vsesoyuznogo Neftyanogo Nauchno-issledovannogo
geolog-razvedochnogo Instituta (VNIGRI), n.s. 85,
227 p., 90 pl.
Sokoloy, B.S. (ed.) (1962). “Osnovy Paleontologu. Tom 2,
Gubki, arkheotsiaty, kishechnopolostnye, chervi.’ 485
p. Izdatel’stvo Akademii Nauk SSSR, Moscow.
Steininger, J. (1831). ‘Bemerkungen iiber die
Versteinerungen, welche in dem Uebergangs-
Kalkgebirge der Eifel gefunden warden.’ 1-44, Trier.
Strusz, D.L. (1966). Spongophyllidae from the Devonian
Garra Formation, New South Wales. Palaeontology
9, 544-598, pl. 85-96.
Talent, J.A. (1963). The Devonian of the Wentworth
and Mitchell Rivers. Geological Survey of Victoria
Memoir 24, 1-118, pl. 1-78. 4
Taylor, M.A. (1984). ‘Geology of the Middle Devonian to
Early Triassic of the Wauchope district, N.S.W.’ B.Sc
Honours thesis, University of Sydney. 158 pp.
Torley, K. (1933). Ueber Endophyllum bowerbanki M.
Ed. U. H. Zeitschrift der deutschen geologischen
Gesellschaft 85, 630-633.
Van Soest, R.M.W. (1984). Deficient Merlia normani
Kirkpatrick, 1908, from the Curagao reefs, with
a discussion on the phylogenetic interpretation of
sclerosponges. Bijdragen tot de Dierkunde 54(2),
211-219.
Webby, B.D., Stearn, C.W., & Zhen, Y.-y. (1993). Lower
Devonian (Pragian — Emsian) stromatoporoids from
Victoria. Proceedings of the Royal Society of Victoria
105(2), 113-185.
Webby, B.D., & Zhen, Y.-y. (1993). Lower Devonian
stromatoporoids from the Jesse Limestone of the
Limekilns area, New South Wales. Alcheringa 17,
327-352.
Webby, B.D., & Zhen, Y.-y. (1997). Silurian and Devonian
clathrodictyids and other stromatoporoids from the
215
DEVONIAN MARINE INVERTEBRATE FOSSILS
Broken River region, north Queensland. A/cheringa
21, 1-56.
Winchell, A.N. (1866). The Grand Traverse Region: A
report on the Geological and Industrial Resources
of the Counties Antrim, Trand Traverse, Benzie, and
Leelanaw in the Lower Peninsula of Michigan. Dr
Chase’s Steam Printing, Ann Arbor. 97 p.
Winchell, A.N. (1867). Stromatoporidae: their structure
and zoological affinites. Proceedings of the Americal
Association for the Advancement of Science 15, 91
-99.
Wright, A.J. (2008). Emsian (Early Devonian) tetracorals
(Cnidaria) from Grattai Creek, New South Wales.
Proceedings of the Linnean Society of New South
Wales 128, 83-96.
Yanet, F.E. (1965). Mikrostrukturnye osobennosti
stenok eifel’skikh 1 zhivetskikh tabulyat 1 khetetid
Urala. In Sokolov, B.S., & Dubatolov, V.N., eds.
Tabulatomorfnye korally devona i karbona SSSR,
Trudy I simposiuma po izucheniya iskopaemykh
korallov 2, 12-24. Nauka, Moscow.
Yu Chang-min & Jell, J.S. (1990). Early Devonian rugose
coral fauna from the Shield Creek Formation, Broken
River Embayment, North Queensland. Memoirs of
the Association of Australasian Palaeontologists 10,
169-209.
Zhen, Yong-yi (1994). Givetian rugose corals from
the northern margin of the Burdekin Basin, north
Queensland. A/cheringa 18, 301-343.
Zhen, Yong-yi (1995). Late Emsian rugose corals of
the Mount Podge area, Burdekin Basin, north
Queensland. Alcheringa 19, 193-234.
Zhen, Yong-yi, & Jell, J.S. (1996). Middle Devonian
rugose corals from the Fanning River Group, north
Queensland, Australia. Palaeontographica A242(1-
3), 15-98.
Ziegler, W., Klapper, G., & Johnson, J.G. (1976).
Redefinition and subdivision of the varcus-Zone
(Conodonts, Midle - ?Upper Devonian) in Europe
and North America. Geologica et Palaeontologica
10, 109-140, pl. 1-4.
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.
DAG,
faabdardoaners ft noe eta instid nahh vate j ; rei
nope ME ders tat ebocaidtonyd shins tik -
pene, elt RAR "Na selina
R all
eure aT hiats (ic seine b, tT, Hy ay EB aba) a
ie tae one
in: ; hi f Bande gent fu idole nee on
Ran ahi
ats
‘Yh tergies setae i eae A oububaiah lege nt
‘aiteniorg’ weal
by toad aii arveneitetotal ls he
Bey 7 ree
“ra ptyet yh be OG DIS f MOVE
YAbtanenionh HERS pops
Heyes aitiagie: tush vache Soa weoweemoeaMeNdi
BL sale! FN eee arstiylh us LA ayabs, te
'. tht y' rai Heal ee a iN eR ARS Oy. |
\ ad Te Diath ‘| fd ipo eh at 1 Viaonieaseata | Coe), 9a Ty
hve Os hbé Arp fitty At Hl FSD ap ats\Aorns
+H washcoainth real hile ie sf3 fatty, nahi el att
" deena i, ‘waren snwhesten beth
‘id yt TE Mt
i
afta ul Beto byte:
SRO lrg wet ie to anebttad, Apmetiar’d orti
bath. -lpeheganiocthl Pil AM a iain bisaelivstentd sith
oat tat Aye cS Hee OMIM sensor anh
r i; wpe un ats tania § CORE Si eas Sivelf i
e . ; inane ot aitt Th Panty pet
a cri hens si sine “itt 7
seals? Wd ponligrea bit enw Saw’ wT ALL
‘Wm det) gnileqse enero!) sdlvd bebagad
a i =y
lave if
sae satan i viol ‘iio pin 1
eon ile sided Jeb “betgatte Tut
is im Wee invotens. ad? a coianrtoy boowstoucll ta Ui
oth a be
alt Pony ey mA: SM sett ai boarrat odw
wipertia ie eed | Thine wif aly oa Oo barton
7 fli bigot Shih itt mat boenet ited ho:
nines Sede arate) teks ri Vitti bel to Tne] eet
(toni OD 25 Fahy ‘ered oor a At
rwautal at soit t0% bao bitty sit alii
wh th suedw A50lH eyaiieall matees only, antral,
rotyrale, fh to Meq modding ot th bazingesst
tort Vey BOE KB TVG St ai: Bh - —
caer te OR,
wih yer: (peel) weal vet inition saoltuse !
galt gundite avitosd oy! & barniieob (200%
yeredire.) oxtl nin iilsool ed 6 han 2p ard
WORE oy OOREIED COURTS AO) baer |
yasotmdT Agoda 00028: 7 bsol ajacit
a) Yio waedaqmmosty bun odie aools oykve:
yt ort ad) Hoggue OW! .oonanta outs to dra
in St thot) owrthon gaibmshcatadlied ite
abe (some!) GORE LEA OORBTD: A ty hoot pesogiee
Feith CURLED hey lad aVh RO A doen die? tow rhitist
“" bsoid NO EE: i belt eu .
ts} chai “yend petto ‘oghndlbin aiden
Dutra pateopurnt 2c 7 Leweepaeo: er sant
ei aes god et bos alin erwok, ons
ht yd ninhove Weotlge na ice |
(a) 0 a aoe
ty como: Bini Pepiaiany cadence A rest
wiht Au ana Namie sid m berate’ Bae G08" 5
han atenalis: ‘bidbnad ey we eit
cai -eceitsea 4a); ee eit
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.
Jia \h
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
Snout & SS —T
\\
1 } \\ xX
NN
Poe NES
>~ 7 (' " \ \
Rees Sm 1 ; \
2 le | Lowe §
Lars Wey \
\ 4-9 | jek ae
Nee A es
. | ae aa
f {ieee S
ok
i=)
el nell cmmastiemsenticennesticemedlienssntieenes tl anmnesthacam dices ’cocenetiteeelimemtianeticnmatiemes tant °,” litecnsesticmae’ianeatienn lt emactinnaticnns!ammatiemmd tensa onmaalicmdicnun!cmentemeeedlmeatiemntieenaltcnctemadimmeticaed
a a as Oe er er ar nr wr car ee ce acer arr ae ae eee ae eee se es
~
I os ee ae |
He teach
| |
\ | \ |
| | , \ |
' 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
p=ulo7sT]
6S #0 TLI
19
— (+6)
[ell LrFLLLi
=(W)
STFS 907
—(9°9)
09 F681
a AO)
[rl ©7F8OLI
— (3°)
[e] S€ FO 191
—(¢L1)
[Le] SEFLIEL
— (79)
[sl] PE FS LSI
—(°2)
l6l eI =8°Stl
—(97@)
[7] STS 9E1
—(F 11)
[gs] 0S FEI
—(0'8)
Or STII
— (Sp)
61 #8 £6
—(r'L)
TF 3°06
= (0)
STF E69
9d
69
%b'9 (L'b1)
lol] +0 F601
%S'°9 (ZS)
SOFS EI
%0°9 (L'9)
POFIII
%b9 (88)
POFLOl
%8°9 (061)
[s] 8066
%TL (11)
[tr] TOF €6
%E9(L 91)
SOFE6
%O'L (9'L)
[6] €0F7OI
%8°9 (L' +1)
[e] L0 +98
%6'L (8'L)
€0F66
%OL(E rT)
9098
MES (p11)
vOFSL
Mrs (ETI)
COFLL
%EOI (S91)
LOFUL
sd
€L
%9 TT (OID)
SOFE OC
%6 IT (rE)
90 9b
%6 OI (61)
70 +007
%0°TI (16)
80 FS 07
%L IT (C6)
LOF68I
%9'EI (S01)
€0 F081
MOLI (LL)
pOoFTsl
%0° ET (78)
SOF E6I
Mp El (CTT)
60F88I
%Y rl (6L)
SOFE SI
%7 ST (8'S)
POFITLI
%6'ST (LL)
SOF6TI
%IST (9'L)
vOFLEL
%OLI (MO
LOFS8TI
pa
€=ul6+T]
COFOFI
79
%eS (P81)
[st] pO 16
MES (STI)
60FT' II
%7S (L'07)
le] T1456
%Ib (TST)
[r] LOFTS
%L'S (C91)
90F8'8
%L"L (S81)
[Le] €0 +86
%79 (9°81)
[L190 +06
%CL (OFT)
[s]SoFrO0l
%9°9 (FST)
L0*66
%9'L (LL1)
SOF16
%9'6 (TET)
[rp] 71 = LOI
%8'OI (S Sz)
[rl € 1 $71
%0'6 (C81)
[L190 E8
%IEl (791)
60F16
ed
89
%9 17 (S01)
[st] VL +T6E
%OIT(S LI)
SSS bb
%6 1 (LL)
[el S1 +L 6¢
%9 17 (8°S)
OLFPLE
%b IT (S01)
[s] 81 ¥9'8€
%79T (6'8)
ler] SOF STE
%ETC (CL)
LOIS OFL +E
%EST (O'L)
SOFTLE
%8'9T (6'L)
irl pL F8'rE
%L LT (8'p)
SOFEVE
%C-8z (E'S)
SOFTTE
%BTE (L' 11)
91 + 8'0E
%6 OE (Z'EL)
[LI r LF 6LZ
%L ve (E01)
VIF be
cd
p=uloet]
90F8'II
89
%0'SI (CTI)
[r1] 60+ £97
%EST (LSI)
SEFSTE
%L rl (FL)
[el 11 + 9°92
%T~SI (Ch)
[rl] $0 +092
%8'+I (9'6)
[s]O1L CH
%6 LI (56)
ler] €0 + 9°E7
%I-SI (901)
lol] 80 #L'€Z
%0'LI (8'L)
90 F6 be
%6 LI (r'€)
[Ir] +0 + O'F~
%9 61 (9'L)
90 ¥ Ibe
%L6I (8')
SOF TC
%S CC (TCI)
T1+1I
%OTT(S TL)
60 +961
bre (E11)
T1L¥691
Id
Si
II
Ol
€
(u)
ezis ojduies
¢=ulose]
OCFETE
el
Ol
mG
(A) o8y
[siayorrg Ul onyeA “xeUT]
WO YNZ < SOTPUI [[v 1OF Uv
18}0],
yNpVy
ynpeqns
SUIpIvO,
dng
dnois os
yal
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
89 €L [BIOL
— 0°98 C el
[RG acOIle 10r -/A9Cll SiS vr -v98I VL ‘L€91 1G Hf V Ol
Pe we bees Spee Wie co [rv] pee ech [rl]
€8-VIIl | vSl-S O€1 £6 :-9791 | LIL -S VLI 19 {Or € 8-106 18-072 ey (EE S 6
Pere OO eo Le] [s] aL [s] [s]
Lot] “ofc 66) pic: [3] 9. “c- ‘9: Lot] lor]
[6] [6l oe ae (6l [6] apes tee er) a
op nee lesisiso com = voneoige | cor ove leo ea I) | Bec cececsl ee ee S| 6
891 -0 £9 86-696 | S CI- -6 07 € 1-9 90--L IV |
. ‘ . . © . . 6 . [3] [2] . 6 . . © . . 4 . . 4 .
Of -S 6E 66:CIL VL-8S9 6 OIG ZA as CVI -18et 99-1 SS | JES) ok, (OY 68 -9 CV 6 14
Gale Cces 9°91 87S 9°67 -€'VS TET VOL €07 :L' C9 SOM LO | OSI lS Cin oll Ce 1S OTE S te J
LS 6 1S OEE OR ell Ol 617: EL VE-ESE 07:76 88 -L'97 TOI 0°87 LL 9% iS ¢ | Wapeqns
EET CEI WAG WILE 667 -6'67 SOE COE 6 0€ -6 0€ il it COI “S91 191 4 PSI of SM 8 [ | Surpieox
- - ; - - ea
(W) | (4) dno
oZIS
(TdS) 94 cd tas ca De | Fee ay o3y
“SISATVUL JY) WIOA S[VUTIUL IUOS JO UOISN[IXO OY} 0} INP 9ZIs s[ dures [v}0} [enbo you svop
Siu) o10ya [u] sjoyov1q a1enbs ul UdAIs ozIs ajduleg -oAQQT x ['’4/(' A —’A)] “ort ‘suMM]OD JULATA1 JY) JO opis puvy JYSII ay} UO pojyussaid aie “(T-} =
av) 1vaXk snorAaid ay) 0) JAQLIAA YIMOIS 107 Sane ‘dnd z sv azIs UMOUY 94} SI QA puL CT ‘OT “6 ‘8 ‘ZL ‘9 ‘S ‘FE ‘ZT =} ELOY %OOT X [ACA — *A)| Corr
‘SUUIN[OD JUVAIII.A JY} JO OPIS puvy Jo] 94} UO pajuasoAd 9.18 0.19Z ISU 0} JATIJLIIA YJMOAG OJ SIN[VA *PIPAOIIA JOU IAA S[RUIUL ISI} JO Z[ 10J STS “9't
‘SUOIIPUOD YSNOA JO asnevd0q TAS P1090. 0} a[qissod sve jou svAd JI (YD}vI-Aq) vas Jv PoANSvoU S[eUTIUL 10,7 ‘s[VUITUL (posdse}) 95v-UMOUY puL pase
-u0l UEP Jo AaquInu ay} SI (U) 9ZIs aJduIKS *(Z B]quUI 99S) SISATeUL WOA, PIpNIXo 919M [[q pue Lg ‘Cg SorqulAVA *7Z 9Iquy, 105 sv (ZL q-Tq) Seyquriva
*‘pojuasaid os[e Si (oSv-UMOUYUN JO SopVUl) o[quIIvA
YIva JO INVA WINUIIXLJA] *S}OYIVAG Ul UOHLLIVA JO JUIIDYJIOD & YJIM “YS F UD Ul 91¥ SJUIWIAINSvOU [TY Soy aeak snoraaid ay) uso. (11) pue $A
WO ‘0.192 98v ye (1) :sjvas ny UVdILAPY YINOS a[vUl Jo USWIOANSvaU ApoOg JO aN[vA ULI JY} 0} VANLIAA (71 G—-1q) Sorquiaea Apog Ul YYMOAD :¢ [Quy
WY,
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
Ee
sce
£
L=))
c
2
>
2)
(s)
a
| wes
|
oF vv ov Or gE gE ve oF vy ov OF ge 9 ve
(w9) Jaddiyjes0) 0} jnous U4 (w3) seddijjas0j 0} jNOUS UT
a4
<0
E < om OO
(=)
2 omoo o L
> <O4 000 o
Oo
> O «e405 o L
8 O fa <@« OO
=
= aa <4 F
<4 <4 |
a4 <4 r
EO)
oO Ci <a
On
Sa ae AS SS Se LT RS ee Sa
cat 0c 82 9¢ ce OF 82 92
(WO) 18a JO S1yU99 0} JNOUS UF (wo) 420 JO 4]U99 0} JNOUS UZ,
E
=
£
f=)
c
ao)
>
co)
fe)
Re)
c
ae
ov ge gE ve ee ge 9€ ve ce
(ud) efe ye peay Jo sduaJaywiNdJIO U7
(wis) aXe ye peasy Jo sousia}WINIIIO U7
Proc. Linn. Soc. N.S.W., 130, 2009
12
10
Age (y)
Age (y)
Age (y)
23k
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
Ln length of hind flipper (cm)
4.2 44
46
Ln body length (cm)
48 5.0 5.2
Ln length of hind flipper (cm)
Age (y)
Figure 5a, b: Bivariate plot of log length of hind flipper (cm) on: (a) log length of seal (cm)
and (b) age (y). PEM animals, open squares; MCM animals, closed triangles.
cm (v = 8). The maximum recorded value for B10
was 55.0 cm (animal PEM1560, SBL 201 cm).
Length of hind flipper (B12)
Growth of length of hind flipper (B12) was rapid
during the early postnatal period and continued to
increase until at least 8—9 y (Table 2 and 3). Overall
growth scaled with negative slope relative to SBL (b=
0.81) and expressed negative allometry relative to age
(Figures 5a, b; Appendix 3, 4), increasing by 103%
at 10 y RTP (Table 3). Growth increment decreased
with increasing SBL until about 4 y (c. 15% of SBL)
(Table 2). The mean B12 of all males > 10 y, including
unaged animals > 200 cm was 28.7 + 0.9 cm (n = 7).
The maximum recorded value was 32.0 cm (animal
PEM1890, SBL 192 cm, = 12 y).
Body length as an indicator of age
In animals 1-10 y, growth in SBL was highly
positively correlated with age (y) (r = 0.96, n = 56)
(Appendix 4). After fitting the (robust) straight line
model of age on standard body length, graphical
displays of residuals and fitted values were examined,
and the straight line model was found to be adequate.
Thus, the following equation can be used as a ‘rough
indicator’ of absolute age for animals 1—10 y.
Age = - 6.54 + 0.0087 x SBL, n = 56
The coefficient of variation (100 x s/X) in SBL for
young males I—5 y (17.2%) was considerably higher
than 1n older males (8-10 y, 6.9%; => 12 y, 5.3%).
ps3)
Body length as an indicator of age group
Linear discriminant analysis was used to classify
seals of unknown age into one of the four age groups
(pup, yearling, subadult, adult) based on body length.
Performing linear discriminant analysis using the
body length data where the age group is known we
get the following four linear discriminant functions
of the form y = mx+b:
VA = OO <SBE 65 (pup)
y, = 0.25 xSBL -11.14 (yearling)
y,; = 0.36xSBL - 23.46 (subadult)
y, = 0.50xSBL - 45.28 (adult)
where, SBL is in cm. A seal with known SBL but
unknown age is classified into the age group which
gives the largest value for the associated linear
discriminant function. For example, an animal 150 cm
long would have linear discriminant function values
OLY = 2205 = 20-308 Vane 30-54 andy te 29) 2 ane
so would be classified as a subadult. Animals over
180 cm would be automatically classified as adults.
Table 4 shows that when the method was used
on animals of known age it was highly successful in
classifying animals into the correct categories. All
3 pups were correctly classified and nearly all the
yearlings (7/8) were correctly classified but one was
classified as a pup. There were some difficulties in
distinguishing yearlings with subadults and subadults
with adults but only one adult out of 22 was incorrectly
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
Table 4: Discriminant analysis for seal age group (pup, yearling, subadult, adult) inferred from body
length of male South African Fur seals.
Size (i) is at age zero, RGR y0; and size (ii) from the previous year, RGR yt-1. All measurements
are in cm. Sample size (n) is the number of seals of known-age (MCM animals tagged as pups), and
aged from counts of incremental lines observed in the dentine of upper canines (PEM animals), n = 70
(of the 73 animals of known age, three animals had insufficient data for this analysis to be carried out).
Includes animals >=12 y (known to be at least 12 y but could not be aged more definitively due to the
limitations of the dentition aging method). Percentage of animals correctly classified into age group is
given in brackets. The overall percentage correctly classified: (3+7+23+21)/70 x 100% = 77.14%. Pups:
All 3 pups have been correctly classified. Yearlings or juveniles: 1 yearling was incorrectly classified as a
pup and the rest of the juveniles (7) have been correctly classified. Subadults: 9 subadults were classified
as yearlings, 23 subadults were correctly classified and 5 subadults were classified as adults. Adults: one
(1) adult was incorrectly classified as being subadult and the rest (n = 21) were correctly classified.
True Group
Juvenile or
Predicted Group Ea Yearling(7 mo to | Salt art geese aie 35) Total
mo) 610) to 7 y 6 mo) 7 mo)
— 3 (100%) 1 (12.5%) 0
Juvenile or F
wens 0 7 (87.5%) 9 (24.3%) 0 16
Subadult 0 0 23 (62.2%) 1 (4.5%) 24
Adult 0 0 5 (13.5%) 21 (95.5%)
Total 3 37 22
classified as a subadult. The overall percentage
correctly classified was calculated from adding up all
the correctly classified animals and then dividing by
the total number of animals multiplied by 100%; that
is, (3+7+23+21)/70 x 100% = 77.14%. Body length
is therefore useful in discriminating between different
age groups but some groups such as yearlings and
subadults can be difficult to correctly classify.
Curvilinear Body length as an indicator of SBL
Curvilinear body length (CBL) was found to be
approximately 10.0 cm longer than SBL (SBL: 146.7
+ 5.6; CBL: 157.1 + 6.2, nm = 50 using paired samples
only). However, CBL was greatly influenced by the
quantity of food in the stomach and by the degree of
post-mortem bloating. For example, CBL was 20-25
cm longer than SBL in 5 animals that had been dead
for several days, or had consumed large quantities of
fish; therefore, CBL was not considered to be a useful
substitute for SBL.
Growth Curve Models
Figure 6 shows a non-linear least squares fit of the
Logistic model (Equation 3) to SBL vs. age for male
Proc. Linn. Soc. N.S.W., 130, 2009
South African fur seals compared to curve fits on data
from a previously published study on the Australian
fur seal (Arnould and Warneke, 2002). Non-linear
fits were also made using the exponential saturation
+ constant or von Bertalanffy model (Equation 2),
and the Gompertz or double exponential equation
(Equation 4). Table 5 shows the statistics of the curve
fits. The correlations for all three models are very
high (r > 0.94). Tests for significant differences in the
fitted parameters were done using t-tests assuming
equal variances or assuming unequal variances as
appropriate (Cochran, 1977).
The Australian fur seal data fits to the von
Bertalanffy model quite well (Table 5), however the
model does not appear to be suitable for the South
African fur seal data. The fit for the South African
fur seal data gives a fitted curve that is very close to
linear and gives an unrealistically high estimate of the
asymptotic SBL of over 270 cm. South African fur
seals have a lower apparent pup size and exponential
constant and the growth rate is lower than for the
Australian fur seals (Table 5).
The fits using the logistic (Equation 3) and the
Gompertz (Equation 4) equations are more similar to
233
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
250
200
150
Standard Body Length (SBL) cm
50
SBL data for Australian fur seals
SBL of South African fur seals
0 5 10 15
Age (y)
Figure 6: Growth kinetics of male South African fur seals (Arctocephalus
pusillus pusillus) (closed circles) compared to male Australian fur seals
(A. pusillus doriferus) (open squares). Age in y and SBL in cm. Curves
fitted using the logistic model (Equation 3).
each other than those made using the von Bertalanffy
model (Equation 2) and both models give more
realistic estimates of asymptotic maximum size for
both the Australian and South African fur seals. Table
5 shows that the significant differences in the model
parameters between the South African and Australian
fur seals are the apparent pup size (P) and the
exponential constant (k). The significant differences
in (k) values reflect a slower growth rate from a lower
initial (P) in South African fur seals.
The asymptotic maximum SBL is about 214-232
cm based on the logistic and Gompertz models. These
two models agree that the asymptotic maximum SBL
is not significantly different in South African and
Australian fur seals. Overall, the logistic curve (Figure
6, Equation 3) seems to be the most satisfactory growth
model, based upon the high correlation coefficients of
least-squares fits to the data and the lowest relative
errors of the fitted parameters.
234
——e« Predicted SBL for Australian fur seals
DISCUSSION
Age determination
Dentition-age estimates of
the South African fur seals were
considered to be reliable, with
inconsistencies among _ readings
mitigated by repeated estimates,
following a set protocol of procedures
and double-blind tests (Payne, 1978,
1979; Doubleday and Bowen, 1980;
Arnbom et al., 1992; McCann, 1993:
Oosthuizen, 1997; Oosthuizen and
Bester, 1997; Arnould and Warmeke,
2002). Nevertheless, the limitations of
dentition-based estimates of the ages
of seals are apparent, particularly for
old animals where the pulp-cavity has
filled and they can no longer be aged
by growth rings. There is a need for
more life-history and morphometric
data based on animals tagged as
pups.
Predicted SBL for South African fur seals
Body size
Arctocephalus pusillus is the
largest of the fur seals with the South
20 African subspecies (4. pusillus
pusillus) tending to be slightly smaller
than the Australian subspecies (A.
pusillus doriferus) (Stewardson et al.,
2008). Comparison of growth curves
for the two populations show that the
Australian fur seal grows at a faster
rate than the South African variety
but asymptotic maximum sizes are very similar
(Table 5). Male SBL ranged from 66 to 243 cm.
The largest animal in the collection (PEM952) was
measured in 1980 at Kings Beach, Port Elizabeth,
by V. Cockcroft and A. Bachelor. This is of similar
length to an unusually large male (SBL 241 cm)
measured by Rand in 1946 (Rand, 1949). The largest
animal measured by the first author was 203 cm in
1994 (PEM2201). The largest individual in the data
set used by Arnould and Warneke (2002) for the
Australian fur seal was a 15 y old bull 224 cm long
(close to the asymptotic maximum sizes estimated
using the Logistic and Gompertz models for the
Australian fur seal). The slightly larger mean size
reached by male Australian vs. the South African
populations of Arctocephalus pusillus may or may
not be genetically based. Stewardson et al. (2008)
pointed out that the present South African population
has largely recovered to pre-exploitation levels,
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
Table 5: Growth Kinetics Models of Male South African fur seals (Arctocephalus pusillus pusillus) com-
pared to Male Australian fur seals (A. pusillus doriferus).
The SBL vs. age data were fitted to the exponential saturation model (Equation 2), the Logistic
model (Equation 3, Figure 6) and the double exponential of Gompertz model (Equation 4). Student’s
t-tests were performed to test if the fitted parameters for the South African and Australian fur seals
were significantly different (Cochran, 1977). Preliminary F-tests showed that in most cases the vari-
ances could be assumed to be equal (p > 0.05). This assumption could not be accepted in the cases of
the asymptotic maximum sizes and incremental growths determined using the exponential saturation
model. The t-test for the case of unequal variances was used for comparing the asymptotic maximum
size and incremental growth of the two varieties of fur seal estimated using the exponential saturation
model. Growth curve fits for the South African fur seal are based on 73 animals of definitive age (denti-
tion-aged animals with an undefined age >12 y are excluded). The data for the Australian seals (n = 69)
were redigitized from Arnould and Warneke (2002).
5a. Exponential saturation growth model
seeea!
Maximum SBL
(cm)
Pup Size at Birth | Lifetime Incremental
Growth (cm)
Population
Australian fur seal
n= 69,
r= 0.9423
-0.1703 139.4
+ 0.0223 ay OD + 8.54
South African fur
seal
n= 73, r=0.9524
-0.0799 VAM 203.5
+ 0.0197 ge 3.117) 32 3.55
P<0.001
Significance P<0.001 P= 0.0678 n.s.
P=0.1928 ns.
5b. Logistic growth model
Maximum SBL Growth k Pup Size at Birth | Lifetime Incremental
(cm) (y') (cm) Growth (cm)
Population
Australian fur seal
n= 69,
r= 0.9443
South African fur 139.6
seal ee + 9.697
+ Q.
n= 73, r= 0.9495 a
Significance jon (OL OATS) tess p < 0.001 p= 0.1796 ns.
5c. Double exponential or Gompertz growth model
-0.3023 124.2
+ 0.0268 ae) // IIS)
Maximum SBL
(cm)
Growth k Pup Size at Birth | Lifetime Incremental
(y") (cm) Growth (cm)
Population
Australian fur seal
n=69,
r= 0.9437
130.2 + 6.569
224.2 -0.2352 94.04
9-138 + 0.0242 a3 uliO7
South African fur
seal
n= 73,r=0.9528
158.3 + 14.99
p = 0.0954 n.s.
Proc. Linn. Soc. N.S.W., 130, 2009 DBS
Misys} -0.1599 74.01
+ 14.48 + 0.0207 + 3.844
Significance P= VOLTS nes: p= 0.019 p < 0.001
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
whereas the Australian population is still rapidly
increasing and have not yet reached a steady
population.
At birth, male South African fur seals are about
35% (c. 70-80 cm) of their mean adult size which is
about 197 cm based upon the mean adult size, SBL
for animals >10 y including unaged animals > 200
cm. At puberty they are about 57% (c. 112.8 cm
at 3 y) of their mean adult size. Although axillary
girth varies with body condition, it is usually about
57-67% of SBL. The foreflippers are relatively long
measuring 25-26% (c. 18 cm) of SBL in pups, and
24% (c. 48 cm) of SBL in adults. The hind flippers
are considerably shorter measuring 19% (c. 13 cm)
of SBL in pups, and 14.5% (c. 29 cm) of SBL in
adults.
Body shape
Male South African fur seals are exceptional
swimmers and divers, and haul out on land to rest,
moult and breed. Body shape and general physiology
have been modified to accommodate the demands of
both marine and terrestrial environments (Bryden,
1972). For example, bulls spend most of their life at
sea, hauling out to moult (predominantly February
and March), rest, and reproduce (establish territories
and breed from late October to late December/early
January).
The body is streamlined with a rounded head
and a relatively short snout; small external ear
pinnae (narrow and pointed); a small tail positioned
between the hind flippers; a retractable penis that can
be withdrawn into a cutaneous pouch; and modified
fore/hind limbs (flippers).
The strong fore limbs have been modified into
elongated flippers for propulsion through the water
(forceful strokes towards the body) and terrestrial
locomotion (palm extends laterally with the flipper
bending between the two rows of carpal bones).
Characteristic features include predigital cartilage, a
long first digit, reduced fifth digit, rudimentary nails
and hairless palms.
Unlike the foreflippers, which are the primary
appendage used for propulsion through the water,
the smaller hind flippers have been modified for
terrestrial locomotion (soles extend laterally with the
flipper bending forward at the ankle). Characteristic
features include predigital cartilage; long grooming
claws on digits 2-4; enlargement of digits one and
five; and hairless soles.
Function and growth
Overall growth in SBL was similar to that of
other highly polygynous male otariids including
Arctocephalus gazella and Callorhinus ursinus, with
236
rapid early postnatal growth; a sudden increase in
body size at puberty; and a reduced rate of growth
soon after attainment of social maturity (McLaren,
198)
South African fur seals pups are born on land
between October and late December (Rand, 1956;
Rand, 1967; Shaughnessy and Best, 1975). Newborn
pups are 70-80 cm long at birth (c. 35% of mean adult
length), which agrees with the apparent pup size
estimates from the present study shown in Table 5.
In November (when the majority of pups are born),
mean length and weight is about 76 cm and 5.986 kg
for males, and 73 cm and 5.487 kg for females (Rand,
1956). By April, mean length and weight is about
82.0 cm and 19.183 kg for males, and 84 cm and
15.147 kg for females (Rand, 1956). Table 5 shows
that the estimated pup size at birth derived from the
exponential and logistic growth curve models are not
significantly different from the actual measurements
given by Rand (1956).
When juveniles gain their permanent teeth
(June) they disperse to deeper water for short periods,
supplementing their milk diet with solids (Rand,
1956). During this period they learn foraging skills
while accompanying their lactating mothers to sea.
Most animals feed independently at 9-11 months
(Rand, 1956). There is a decline in body weight soon
after weaning (Rand, 1956).
Most males attain puberty between 3-4 y, as
evidenced by the presence of sperm in the epididymis
of some animals at 2 y 10 months (Stewardson et al.,
1998). The onset of puberty (2—3 y) is associated with
a sudden increase in body size (present study). It is
thought that puberty is attained when seals reach a
certain threshold size in body weight, with slower-
growing animals reaching puberty later than faster-
growing animals (Laws and Sinha, 1993). Although
pubertal males produce sperm, they do not have the
ability to acquire and maintain a harem (Stewardson
et al., 1998). Small body size and inexperience
prevents young males from gaining the high social
status required for a breeding male.
Growth in SBL continues to increase steadily
until about 6 y. In animals = 7 y, growth continues to
increase but at a slower rate (Tables 5 and 6, Figure
6). Social maturity is attained at about 9-10 y and
appears to be associated with a weak secondary
growth spurt in body size (present study). At this age,
large body size has a direct advantage in competitive
interactions with rival males, including intimidatory
display without actual fighting, and an indirect effect
through the presence of large stores of fat which
enable large males to remain on territory for up to
40 days (Rand, 1967; Wartzok, 1991). Successful
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
bulls may hold harems multiple times over a two to
three year period but are likely to die before reaching
reproductive senescence (see Stewardson et al., 1998).
Growth in body size slows soon after attainment of
social maturity (present study).
Growth of length of the foreflippers continued to
increase until at least 13 y, with a significant increase
in length at 4-5 y (present study). This increase may
partially reflect changes in swimming and/or diving
behaviour, with older animals presumably diving to
deeper depths in search of prey. Growth of the smaller
hind flippers slowed much earlier (8—9 y) than growth
of the foreflippers (as is also found in the case of the
Australian fur seal; Arnould and Warneke, 2002). The
maximum sizes of the fore and hind flippers found
in the present study for the South African fur seal
are similar to the asymptotic sizes found in the fore
and hind flippers of the Australian fur seal (Arnould
and Warneke, 2002). No special development of
the foreflippers or hind flippers associated with
locomotion was reported in Arctocephalus gazella,
i.e., a more or less constant rate of growth from age
one to 7 (Payne, 1979).
Body length as an indicator of age
SBL could not be used reliably to assign a seal toa
particular age because there was considerable overlap
between year classes, especially among middle-aged
animals. Similar findings have been reported in other
species of pinnipeds (e.g., Laws, 1953; Bryden, 1972;
Bengston and Sniff, 1981). However, SBL was found
to be a ‘rough indicator’ of age for animals 1—10 y,
and of age group (Table 4). The curvilinear models
(von Bertalanffy, logistic and Gompertz models)
shown in Table 5 could also be used to estimate age
from SBL but inspection of Figure 6 clearly shows
that they would not be reliable for estimating the age
of animals greater than about 10 y.
In male South African fur seals, postnatal growth
is rapid with a significant growth spurt at the onset
of puberty (2-3 y) and a weak growth spurt at social
maturity (9-10 y). Body size continues to increase but
at a slower rate between 6 and 7 y, and then growth
slows soon after the attainment of social maturity.
Growth was a differential process and not simply an
enlargement of overall size. 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 with negative slope or were
negatively allometric. SBL was found to be a ‘rough
indicator’ of age and of age group.
Proc. Linn. Soc. N.S.W., 130, 2009
Model Growth Curves for Male South African
and Australian fur seals
Further information is needed on older animals
of known-age in order to more accurately estimate
asymptotic maximum size (see Figure 6 and Table 5).
In the present study, low sample size at the intermediate
ages, and the absence of very old animals of known-
age (15-20 y), made it difficult to determine a more
exact shape of the growth curve. Published growth
curves are also available on the male Australian fur
seal (Arnould and Warneke, 2002, n = 69), male and
female New Zealand fur seals (Dickie and Dawson
2003, males n = 64), male New Zealand fur seals
(McKenzie et al., 2007, n= 86), subantarctic fur seals
(Bester and Van Jaarsveld, 1994) and the male Steller
sea lion (Winship et al., 2001, n = 203). The breeding/
non-breeding status of the animals in the present study
was not known. Breeding bulls are thought to be
larger in size than non-breeding bulls of the same age;
therefore, the growth pattern of male fur seals may be
more complex than implied by the models used in the
present study. For example, the data of Arnould and
Warneke (2002) is based on males shot at a breeding
colony and so has many large males of breeding
status. The males in the present study are mainly
based on dead or dying animals found stranded on
the coastline and incidental drownings from trawling.
The study area of the present study was a seal feeding
area rather than a breeding colony.
Figure 6 and Table 5 show that the von
Bertalanffy, Logistic and Gompertz models suggest
that the kinetics of growth in SBL vs. age is different
in the two subspecies. South African fur seals seem
to have a smaller apparent pup size and a slower
growth rate than the Australian fur seal once living
independently. The skulls of male Australian fur
seals are significantly larger than South African
male seal skulls (Stewardson et al., 2008), however
the often repeated statement that Australian fur seals
are consistently larger in body size than the South
African variety (Pemperton et al., 1993; Arnould
and Warneke, 2002; Stewardson et al., 2008) is not
supported by the values for the asymptotic maxima
of the logistic and Gompertz curve fits shown in
Table 5 and by inspection of Figure 6. However, for
our South African material, the average SBL of all
males > 10 y and/or SBL > 200 cm was 197 + 4.1
cm (n = 15) is significantly smaller than a similar
calculation for Australian males (211 + 1.5 cm, n =
17) using the data of Arnould and Warneke (2002).
This might more accurately reflect differences in the
types of populations sampled in the study by Arnould
and Warneke (2002) — a breeding colony, and in the
present study — a feeding population probably with
DB]
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
many non-breeding males. In any case, more data on
age-tagged old males is needed to better define the
growth kinetics of the species.
After weaning, the Australian variety seems
to grow faster and reaches maturity earlier than
the South African variety. For example, the data of
Arnould and Warneke (2002) includes one individual
that exceeded 200 cm when only 5 and a half years
old.
Such growth kinetics are consistent with what
would be expected of a rapidly increasing population,
not limited by natural resources, recovering from
severe depletion. The Australian population has not
yet reached pre-exploitation population size whereas
the South African population is today close to pre-
exploitation levels (Pemperton et al., 1993; Arnould
and Warneke, 2002). Differences between the two
varieties might therefore reflect differences between
a well-fed expanding population and a population
in steady-state with limited resources (Stewardson
et al., 2008), rather than a genetic difference. Such
a proposition would predict that as the Australian fur
seal population approaches the carrying capacity of
its niche, a reduction in the average pup size, growth
rate of the pups and the growth rate of independent
animals would be expected.
Winship et al. (2001) working with a data set of
203 aged male Steller sea lions were also able to show
that the Logistic and Gompertz models were better
(in terms of sum of squares residuals and correlation
r) than the von Bertalanffy model for describing the
growth kinetics of seals. The logistic and Gompertz
models are also a very good fit to the growth kinetics
of male New Zealand fur seals (Dickie and Dawson
2003; McKenzie et al., 2007). The Logistic and
Gompertz exponential constants found for the New
Zealand fur seal are comparable to those found in
the Australian fur seal due to a similar lifespan and
similar relative sizes of the pup to the adult.
A great deal of effort has been spent in discussing
the relative merits of growth curves in biology but
often the data sets are too small for this time and effort
to be justifiable (Zeide, 1993). The von Bertalanffy
model was not a satisfactory fit for the South African
fur seal data because it gave an unrealistically high
estimate of the asymptotic maximum size of 276 cm,
which is well above the largest recorded SBL of 241
cm for a South African fur seal. The fitted equation
shows very little curvature, due to a lack of accurately
aged very old animals (Figure 6). Previous attempts
to model the growth kinetics of seals have generally
reached the conclusion that the von Bertalanffy
model tends to give imprecise overestimates of the
asymptotic maximum size (Zullinger et al., 1984;
238
Trites and Bigg, 1992; Bester and Van Jaarsveld,
1994; Winship et al., 2001; Arnould and Warneke,
2002; McKenzie et al., 2007).
The logistic and Gompertz models are both more
realistic than the von Bertalanffy model for post-natal
mammalian growth because they both have a point of
inflection: mammals grow exponentially while pups
and juveniles, then linearly as a subadult and finally
growth decreases asymptotically when reaching
maturity. Both are very popular for modelling growth
in mammals but Zullinger et al. (1984) points out that
bothmodels tend to bothunderestimateandimprecisely
estimate the maximum body size of mammals. Plots
of the fit to the Gompertz model are almost identical
to those made using the Logistic model but Table 5
and Figure 6 show that the asymptotic body size is
not as precisely defined as in the case of the Logistic
model (Equation 4). This is particularly the case for
the South African fur seal data, due to the lack of very
old animals in the data set.
The growth models suggest there are significant
differences in how the maximum size is achieved in
the two populations; independent juvenile and adult
Australian fur seals have a very significantly higher
growth rate than in the case of South African fur
seals. Thus the Australian fur seal achieves maximum
size earlier than the South African fur seal and tends
to grow larger, at least under current population
densities.
Caution is needed in interpreting changes in
growth kinetics of seals over time or differences
between different species or populations. The
population history of the Northern fur seal is similar
to that of most fur seals: extreme depletion by the
beginning of the 20" century, followed by recovery but
for reasons that are not clear-cut population numbers
and growth kinetics have varied considerably since
their initial recovery in about 1940. Trites and Bigg
(1992, 1996) have found that the growth kinetics of
the Northern fur seal (Callorhinus ursinus) has varied
over time on the Pribilof Islands off Alaska but were
cautious about attributing it to changes in population
pressure on resources or other environmental effects.
Trites and Bigg (1992) state that higher growth rates
and body size seem to correlate with lower total
densities of animals but migration effects between
different colonies could be a complicating factor.
Conclusion
The classification criteria for age and age group
developed in this study will be particularly useful
when canines are not available for age determination,
e.g. behavioural studies, census counts and where
animals are drugged for mark/recapture studies.
Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
Removing postcanines for aging live animals as done
by Payne (1978, 1979) might not be possible under
some jurisdictions. Tagging of live animals, should
be encouraged wherever possible, because of the
lack of data on development and longevity of most
species of fur seal impacts on the development of
rational management policies. Information presented
in this study contributes to earlier descriptions of the
South African fur seal (Rand, 1956), and provides
new information on body growth according to age (y)
that 1s useful for comparisons with the Australian fur
seal.
ACKNOWLEDGEMENTS
We wish to express our sincere appreciation to the
following persons and organizations for assistance with
this study: Dr V. Cockcroft (Port Elizabeth Museum), Dr
J. Hanks (WWE-South Africa) and Prof. A. Cockburn
(Australian National University) for financial and logistic
support; Mr B. Rose (Oosterlig Visserye, Port Elizabeth)
who enabled us to collect seals from his commercial fishing
vessels; Dr J.H.M. David (Marine Coastal Management,
Cape Town) for use of measurements from known-age
animals; Mr H. Oosthuizen (MCM) for assistance with
age determination; Mr S. Swanson (MCM) for assistance
with data extraction of MCM specimens; Mr N. Minch
(Australian National University) for photographic editing;
Dr C. Groves for constructive comments on an earlier draft
of this manuscript. This paper was compiled on behalf of
the World Wild Fund for Nature - South Africa (project ZA
- 348, part 1a). The South African fur seal data presented
in the paper is abstracted from a PhD thesis presented to
the Australian National University by C. L Stewardson in
2001.
REFERENCES
Anderson, T.W. (1984). An Introduction to multivariate
statistical analysis, 2nd ed. (John Wiley and Sons
Publ., New York).
Arnbom, T.A., Lunn, N.J., Boyd, I.L. and Barton, T.
(1992). Aging live Antarctic fur seals and southern
elephant seals. Marine Mammal Science 8, 37 — 43.
Amould, J.P.Y. and Warneke, R.M. (2002). Growth and
condition in Australian fur seals (Arctocephalus
pusillus doriferus) (Carnivora: Pinnipedia).
Australian Journal of Zoology 50, 53-66.
Bengston, J.L. and Sniff, D.B. (1981). Reproductive
aspects of female crabeater seals (Lobodon
carinophagus) along the Antarctic Peninsula.
Canadian Journal of Zoology 59, 92-102.
Bester M.N. and Van Jaarsveld A.S. (1994). Sex specific
and latitudinal variance in postnatal growth of the
Proc. Linn. Soc. N.S.W., 130, 2009
subantarctic fur seal (Arctocephalus tropicalis)
Canadian Journal of Zoology 72, 1126-1133.
Bigg, M.A. (1979). Preliminary comments on body length
in female northern fur seal. In ‘Preliminary Analysis
of Pelagic Fur Seal Data Collected by the United
States and Canada during 1858-1974’. Report
to 22nd Annual Meeting of the North Pacific Fur
Seal Commission, pp. 171-179. (National Marine
Fisheries Service, NOAA, Seattle, Washington).
Bychkoy, V.A. (1971). Age variation of body length and
weight of fur seals of Seal Island. Ekologiva 1971
(1), 101-105. [Engl. transl. Soviet Journal of Ecology
2, 81-83.]
Bryden, M.M. (1972). Growth and development of
marine mammals. In “Functional anatomy of marine
mammals’, vol. 1, (Ed. Harrison, R.J.), pp. 2-79.
(Academic Press Publ., London).
Calkins, D. and Pitcher, K.W. (1983). Population
assessment, ecology and trophic relationships
of Steller sea lions in the Gulf of Alaska. In
‘Environmental Assessment Program’. U.S.
Department of Commerce, NOAA, 19, pp. 445-546.
Cochran, W.G. (1977). Sampling techniques, 3rd ed. (John
Wiley and Sons Publ., New York).
Committee on Marine Mammals (1967). Standard
variables of seals. Journal of Mammalogy 48,
459-462.
Dickie G.S. and Dawson S.M. (2003). Age and reproduction
in New Zealand fur seals Marine Mammal Science 19,
173 — 185.
Doubleday, W.G. and Bowen, W.D. (1980).
Inconsistencies in reading the ages of harp
seal (Pagophilus groenlandicus) teeth, their
consequences, and a means of resulting biases. North
Atlantic Fisheries Organization, Scientific Council
Research Document 80/X1I/160, Serial No. N247.
Fiscus, C.H. (1961). Growth in the Steller sea lion.
Journal of Mammalogy 42, 218-223.
Gibbons, J.D. and Chakraborti, S. (1992). Nonparametric
statistical inference, 3rd ed., (Marcel Dekker Inc
Publ., New York).
Johnson, M.L. and Faunt, L.M. (1992). Parameter
estimation by least squares methods. Methods in
Enzymology 210, 1-37.
Krylov, V.I. and Popov, L.A. (1980). On comparative
morphology of Antarctic seals of King George Island.
In ‘Morskie Mlekopitayushchie,’ pp. 40-51. Moscow:
VNIRO [In Russian. ]
Lander, R.H. (1979). Fur seal growth. In “Preliminary
analysis of pelagic fur seal data collected by the
United States and Canada during 1958-1974.
Report to 22nd Annual Meeting North Pacific Fur
seal Commission, pp. 143-170. Seattle, Washington:
National Marine Fisheries Service, NOAA.
Laws, R.M. (1953). The elephant seal (Mirounga
leonina Linn.). 1. Growth and age. Falkland Islands
Dependencies Survey Scientific Reports 8, 1-62.
Laws, R.M. and Sinha, A.A. (1993). Reproduction. In
‘Antarctic seals research methods and techniques’
IB9
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
(Ed. Laws, R.M.), pp. 228-267. Cambridge
University Press Publ., London).
Loughlin, T.R. and Nelson, R. (1986). Incidental mortality
of northern sea lions (Eumetopias jubatus) in
Shelikof Strait, Alaska. Marine Mammal Science 2,
14-33.
McCann, T.S. (1993). Age determination. In “Antarctic
seals, research methods and techniques’ (Ed. Laws,
R.M.), pp. 199-227. (Cambridge University Press
Publ., London).
McKenzie, J., Page, B., Goldsworthy, S.D. and Hindell,
M.A. (2007). Growth strategies of New Zealand fur
seals in southern Australia. Journal of Zoology 272,
377 — 389.
McLaren, I.A. (1993). Growth in pinnipeds. Biological
Review 79, 1-79.
Oosthuizen, W.H. (1997). Evaluation of an effective
method to estimate age of Cape fur seals using
ground tooth sections. Marine Mammal Science 13,
683-693.
Oosthuizen, W.H. and Bester, M.N. (1997). Comparison of
age determination techniques for known-age Cape fur
seals. South African Journal of Zoology 32, 106-111.
Payne, M.R. (1978). Population size and age
determination in the Antarctic Fur seal Arctocephalus
gazella. Mammal Review 8, 67 — 73.
Payne, M.R. (1979). Growth in the Antarctic fur seal
Arctocephalus gazella. Journal of Zoology (London)
187, 1-20.
Pemperton D., Kirkwood, R, Gales R. and Renouf, D.
(1993). Size and shape of male Australian fur seals,
Arctocephalus pusillus doriferus. Marine Mammal
Science 9, 99 — 103.
Rand, R.W. (1949). Studies on the Cape fur seal
Arctocephalus pusillus pusillus 3. Age grouping
in the male. Progress report submitted November
1949, Government Guano Islands Administration,
Department of Agriculture, Union of South Africa.
Rand, R.W. (1956). The Cape fur seal Arctocephalus
pusillus pusillus (Schreber): its general characteristics
and moult. Sea Fisheries Research Institute
Investigational Report, South Africa 21, 1-52.
Rand, R.W. (1967). The Cape fur seal Arctocephalus
pusillus pusillus 3. General behaviour on land and at
sea. Sea Fisheries Research Institute Investigational
Report, South Africa 60, 1-39.
Rosas, F.C., Haimovici, M. and Pinedo, M.C. (1993). Age
and growth of the South American sea lion, Ofaria
filavescens (Shaw, 1800), in Southern Brazil. Journal
of Mammalogy 74(1), 141-147.
Scheffer, V.B. and Wilke, F. (1953). Relative growth in the
northern fur seal. Growth 17, 129-145.
Shaughnessy, P.D. and Best, P.B. (1975). The pupping
season of the Cape fur seal, Arctocephalus pusillus
pusillus. Sea Fisheries Branch South Africa, pp. 1-8,
1975. (Unpublished internal report)
Stewardson, C.L. (2001). “Biology and conservation of
the Cape (South African) fur seal Arctocephalus
pusillus pusillus (Pinnipedia: Otariidae) from the
Eastern Cape Coast of South Africa’, Thesis (Ph.D.),
240
Australian National University. http://thesis.anu.edu.
au/public/adt-ANU20030124.162757/index.html
Stewardson, C.L., Bester, M.N. and Oosthuizen, W.H.
(1998). Reproduction in the male Cape fur seal
Arctocephalus pusillus pusillus: age at puberty
and annual cycle of the testis. Journal of Zoology
(London) 246, 63-74.
Stewardson, C.L., Prvan, T., Meyer, M.A. and Ritchie,
R.J. (2008). Age determination and growth in the
male South African Fur Seal Arctocephalus pusillus
pusillus (Pinnipedia: Otariidae) based upon skull
material. Proceedings of the Linnean Society of NSW
129, 207-252.
Thorsteinson, F.V. and Lensink, C.L. (1962). Biological
observations of Steller sea lions taken during
an experimental harvest. Journal of Wildlife
Management 26, 353-359.
Trites, A.W. and Bigg, M.A. (1992). Changes in body
growth of northern fur seals from 1958 to 1974:
density effects or changes in the ecosystem?
Fisheries Oceanography 1, 127-136.
Trites, A.W. and Bigg, M.A. (1996). Physical growth of
northern fur seals (Callorhinus ursinus): seasonal
fluctuations and migratory influences. Journal of
Zoology (London) 238, 459-482.
Wartzok, D. (1991). Physiology of behaviour in pinnipeds.
In “Behaviour of pinnipeds’ (Ed. Renouf, D.), pp.
236-299. (Chapman and Hall Publ., London).
Weisberg, S. (1985). Applied linear regression, 2nd ed.,
(John Wiley and Sons Publ., New York).
Wickens, P.A. (1993). Life expectancy of fur seals with
special reference to the South African (Cape) fur seal.
South African Journal of Wildlife Research 23, 101
— 106.
Winship, A.J., Trites, A.W. and Calkins, D.G. (2001).
Growth in body size of the Steller sea lion
(Eumetopias jubatus). Journal of Mammalogy 82,
500-519.
Wynen, L.P., Goldsworthy, S.D., Insley, S.J., Adams,
M., Bickham, J.W., Francis, J., Gallo, J.P., Hoelzel,
A.R., Mailuf, P., White, R.W.G. and Slade, R.
(2001). Phylogenetic Relationships within the Eared
Seals (Otariidae: Carnivora): Implications for the
Historical Biogeography of the Family. Molecular
Phylogenetics and Evolution 21, 270-284.
Zeide, B. (1993). Analysis of growth equations. Forest
Science 39, 594-616.
Zullinger, E.M., Ricklefs, R.E., Redford K.H and
Mace G.M. (1984). Fitting sigmoidal equations to
mammalian growth curves. Journal of Mammalogy
65, 607-636.
Proc. Linn. Soc. N.S.W., 130, 2009
APPENDIX 1
South African fur seals (n = 149) examined in this study. Animals were collected from
the coast of southern Africa between August 1978 and September 1997.
Accession Numbers of Specimens used in the Present Study.
Port Elizabeth Museum (PEM), Port Elizabeth, South Africa
PEM603
PEM676
PEM877
PEM928
PEM1159
PEM1698
PEM1895
PEM2007
PEM2020
PEM2049
PEM2087
PEM2186
PEM2203
PEM2257
PEM2401
PEM2414
PEM605
PEM824
PEM886
PEM951
PEM1453
PEM1706
PEM1999
PEM2008
PEM2021
PEM2051
PEM2131
PEM2188
PEM2238
PEM2257
PEM2403
PEM2415
PEM607
PEM828
PEM888
PEM952a
PEM1507
PEM1879
PEM2000
PEM2009
PEM2036
PEM2052
PEM2132
PEM2191
PEM2248
PEM2348
PEM2404
PEM2454
PEM608
PEM834
PEM889
PEM958
PEM1560
PEM1882
PEM2002
PEM2010
PEM2045
PEM2053
PEM2137
PEM2194
PEM2252
PEM2359
PEM2405
PEM2455
PEM658
PEM852
PEM898
PEM975
PEM1587
PEM1885
PEM2003
PEM2013
PEM2046
PEM2054
PEM2140
PEM2197
PEM2253
PEM2374
PEM2406
PEM2458
PEM661
PEM874
PEM916
PEM 1073
PEM1696
PEM1890
PEM2004
PEM2014
PEM2047
PEM2081
PEM2141
PEM2198
PEM2254
PEM2379
PEM2409
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
PEM670
PEM875
PEM917
PEM1135
PEM1697
PEM1892
PEM2006
PEM2015
PEM2048
PEM2082
PEM2143
PEM2201
PEM2256
PEM2400
PEM2411
Marine and Coastal Management (MCM), Dept of Environment Affairs and Tourism,
Cape Town, South Africa
MCM1565
MCM3586
MCM4577
MCM4989
MCMS5001
MCMS5135
Proc. Linn. Soc. N.S.W., 130, 2009
MCM1786
MCM3587
MCM4584
MCM4991
MCMS5002
MCMS5136
MCM2763
PEM3589
MCM4585
MCM4992
MCMS5005
MCMS5142
MCM2795
MCM3636
MCM4595
MCM4996
MCMS021
MCMS5145
MCM3017
MCM4023
MCM4597
MCM4998
MCMS5022
MCM3125
MCM4365
MCM4985
MCM4999
MCM5133
MCM3582
MCM4388
MCM4987
MCM5000
MCMS5 134
241
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
APPENDIX 2
Spearman rank-order correlation coefficients for log body variables of male South African Fur seals.
Variables are as for Appendix 1. Pups were excluded from the analysis. p < 0.001 unless otherwise stated in
square brackets. * Significant at 2% level (2-tailed). ** Significant at 1% (2-tailed).
Sample size (n) in brackets.
BI B2 B3 B4 BS B6 B7 BS B9 B10 Bil B12
Bl 1 082* 0.12 0.74* 0.63 0.77* 0.84% 0.76 0.71* 0.71* 0.82* 0.72*
-99 “OR OO si -97 -94 aSF -54 -96 -98 -96 81 -93
-85
B2 0.82* 1 0.17 0.76* 0.61* 0.62* O.81* 0.78% 0.73% 0.74% 0.86 —0.72*
“986 5102) [OAM Awe LOO -97 -90 -57 Oy <il@) -99 -83 -96
-87
BS 202s ouaON (MODS PS IS O02 OOS TON WOIR MOOT O.08
[0.27] [0.11] -101 [0.02] [0.02] [0.17] [0.99] [0.46] [0.10] [0.26] [0.54] [0.16]
-85 -87 -93 -89 -87 -54 -90 -92 -90 Ti -87
B4 0.74" = 0.76* = 0.25** 1 0.85* 0.84* 0.68* 0.79* 0.74* 0.85% 0.79* 0.76*
070 aes 00) Ne [O!02 eee OS ane 104 -93 Folly | 103s 106 aan alod =85 SV S=ON
-93
BS 0.63* 0.61* ~—0.25** ~—0.85* 1 0.78" 0.68" 0.69* 0.68* 0.72* 0.71% 0.68*
-94 LO 10102] ae 04 eeeT05 -94 57] a pCO peel0S e102 -86 .» 101
-89
B6 0.77* 0.82 0.15 0.84* 0.78* 1 0.96" 0.99* 0.93* 0.92* 0.94* 0.90*
-87 90 [0.17] -93 Oi IB 5] -94 -95 -93 -86 -92
537
B7 0.84* 0.81* 0.002 0.68* 0.68* 0.96* 1 097* 0.92* 0.74* 0.92* 0.82*
-54 Si -54 -61 -57 51 -65 -60 -61 -59 -45 -57
BS 0.76* 0.78* 0.08 0.79% 0.69% 0.99% 0.97* 1 0.93* 0.89* 0.94% 0.90*
-96 Zo OAGI 210308 =100 -94 {50 ul c= ae OY ta nln (09) -84 -99
-90
Bo 0.71* 0.73* 0.17 0.74* 0.68% 0.93* 0.92* 0.93* 1 0.82* 0.89* 0.91%
Hoge STON [OO] =106n 103 -95 -61 Hoa © S0ow E05 ag7 Dr Wto2
-92
B10 0.71* 0.74 0.12 0.85* 0.72 0.92* 0.74" 0.89 —0.82# 1 0.88 0.87%
-96 O) OA =O =i -93 9) i SY =85) ELON
-90
Bll 0.82 0.86 0.07 0.79* 0.71* 0.94* 0.92* 0.94* 0.89% 0.88* 1 0.85%
81 -83 [0.54] -85 -86 -86 -45 -84 OT -85 7 -86
ail
Bl2 0.72* 0.72* 0.15 0.76 0.68% 0.90% 0.82 0.90* 0.91* 0.87* 0.85* 1
-93 206) S[ONIGIN ETON ETON -92 5] £90) 102 elon =36)0 103
Si
Total 99 102 101 108 105 131 65 107 109 107 87 103
242 Proc. Linn. Soc. N.S.W., 130, 2009
C.L. STEWARDSON, T. PRVAN, M.A. MEYER AND R.J. RITCHIE
OTT [B30],
VN VN VN (100>49)060 400+180 61:0 + 16°0- 76 (Za) soddry pury fo yysu0T
VN VN VN (100>49)7%60 4004680 810 = 160- £6 (01a) Joddryor0; Jo ysusT
Looo=4 €61<9:'H (100>49) £60 SOOFITT 770 F EE I- 66 (6) Joddiyor0y oy} Jo UONTOSUT JOLIO}UR 0} jnoUS Jo dry
VN VN VN (100>%)660 100401 L0'0 + SE0- 16 (gq) sutuodo [ey1U08 0} nous Jo diy,
VN WN VN (100>%8L0 4004690 7C'0 ¥ 780" 6 (Sq) odes Jo a[S3ue 0} ynous Jo diy,
1l00>d 16 1>49:'H (100>4) P80 €00FES0 p10 ¥0€0 €6 (pq) Iwo JO x99 0} nous Fo diy,
— - —su (91°) =) S10 — - L8 (¢€q) 249 JO OUD 0} JnoUS Jo dry,
loo>d 988 1>49:'H (100>4)780 10:0 +050 810 + 601 06 (Zq) 240 Je peoy JO soueJEFUIMIAT
10>4 Sg 1>4:H (100>DLL0 +100FLr0 810 + 680 L8 (Tq) ourueo ye peoy Jo soUorEFUIMIIID
(d) Apiqeqorg ~P, Se CO) Same dOIS: els ao an ers] er g]qeliva quepuedaq
AAJOWIOTLV UOISSIAGIA AVIUT'T
(Z 9[QUI, UI So}OUO0OF 998) sISATeUR WOLF popnjoxe [][ gq pue Lg so]qeueA “o[qeordde you 4soq “arr “jour
JOU 919M (q) aUT] dy} Jo odoys oy} jnoge sosoyjodAy jso} 0} posnber suonduinsse jopoul “WN “‘WLOYIUSIS JOU UOISSOIdOI ILOUT] “JOAQMOY “JOU OIOM E_ I[GUIIBA IOJ
suondunsse Jopoyw ‘¢gq wos yede (poyie}-Z) [OAd] %] OU} Je JULOIUTIS ov SUOTLIOIIOD |] YW JUSIOYJoos UONRIEU09 Jop1o-yues uewtvedg “I (Q]] = U “ol “papso0s
-91 JOU dJOM sa[eUl poSeun/pose ¢] Wo sT|gs pue ‘sishyeue wo popnyoxe o19M sdnd) poproser TqS YIM sjeultue poseun pue pose Oj si (U) ozIs o[dureg
*s[voS IN| ULILAPY YING [vu Jo (WD) YSUaT Apog
[ves So] uo (Wd) yUdWOANSvaW Apog So] 10 AjoWOT]e pue ‘s}Ud1DYJo09 UOHL[IA109 19p10-yuRA ULULIvads ‘suoyEenbea aur yy sI¥.48 sarenbs jsvoy .Jsnqoy,
€ XIGNHddV
243
Proc. Linn. Soc. N.S.W., 130, 2009
BODY MEASUREMENTS OF SOUTH AFRICAN FUR SEALS
APPENDIX 4
‘Robust’ least squares straight-line equations, Spearman rank-order correlation coefficients, and allometry for log body measurement (cm) on age
(y) of male South African Fur seals.
Sample size (n) is the number of skulls with body variable and age recorded (only animals 1—10 y were included in analysis, 1.e., n = 68). r, Spearman
rank-order correlation coefficient. All correlations are significant at the 1% level (2-tailed), except for B3. Model assumptions were met for variable B3;
however, linear regression not significant. NA, model assumptions required to test hypotheses about the slope of the line (b) were not met, 1.e., test not
applicable. Variables B7 and B11 excluded form analysis (see footnotes in Table 2).
Dependent variable (B)
Circumference of head at canine (B1)
Circumference of head at eye (B2)
Tip of snout to centre of eye (B3)
Tip of snout to centre of ear (B4)
Tip of snout to angle of gape (B5)
Standard body length (B6)
Tip of snout to genital opening (B8)
Tip of snout to anterior insertion
of the foreflipper (B9)
Length of foreflipper (B10)
Length of hind flipper (B12)
Total
Sample size
(n)
63
63
Sil
68
64
56
67
68
67
64
68
Intercept + S.E.
-2.59 + 0.50
-2.63 + 0.43
2.67 + 0.02
2.03 + 0.03
4.45 + 0.02
-1.28+0.14
3.56 + 0.03
3.10 + 0.03
2.64 + 0.02
Linear regression
Slope + S.E.
0.17 + 0.021
0.12+0.01
0.04 + 0.004
0.04 + 0.005
0.08 + 0.003
0.02 + 0.001
0.10 + 0.005
0.07 + 0.005
0.07 + 0.004
nO) Alternative
P hypothesis
0.59 (p < 0.01) IH 2 [Bl
0.69 (p < 0.01) NA
-0.008 (p = 0.95) ns
0.69 (p < 0.01) NA
0.56 (p < 0.01) 1s re Sa
0.96 (p < 0.01) NA
0.93 (p<0.01) NA
0.90(p<0.01) H,:B<1
0.82 (p<0.01) NA
0.93(p<0.01) H,:B<1
Allometry
df
61
NA
66
NA
62
Probability
(p)
p<0.01
NA
p< 0.01
NA
p < 0.01
Proc. Linn. Soc. N.S.W., 130, 2009
244
Linnaeus: King of Natural History
PauL ADAM! AND ELIZABETH May?
' School of Biological, Earth and Environmental Sciences, University of NSW, Kensington, NSW, 2052;
* School of Biological Sciences, University of Sydney, NSW, 2006
Adam, P. and May, E. (2009). Linnaeus: king of natural history. Proceedings of the Linnean Society of
New South Wales 130, 245-250.
Linnaeus’ legacy was far more encompassing than taxonomic. We argue that, while the systematic recording
of species remains fundamental to modern ecological concerns, Linnaeus also laid the foundation for
other major areas of ecology, including comparative biogeography, plant demography, and comparative
anatomy.
Manuscript received 18 October 2008, accepted for publication 21 January 2009.
KEYWORDS: Biodiversity, Linnaeus, natural history, natural philosophy, taxonomy.
INTRODUCTION
Linnaeus is one of the towering figures in the
history of biological science. He is remembered today
chiefly for his introduction of the binomial system of
nomenclature and for his taxonomy. However, if we
consider his skill as an observer of the relationships
between the plants and animals he classified, and
their environment, he should also be regarded as
one of the earliest practicing ecologists. He passed
on these skills to his many students, including those
who travelled the world collecting and classifying
organisms and became known as the ‘apostles’.
In eighteenth-century Sweden, Linnaeus was
accorded high status, and this great respect continued
to hold internationally until the twentieth century.
Lord Rutherford’s throw-away comment that ‘(a)ll
science 1s either physics or stamp collecting’ reflected
the marginalization of taxonomy and natural history
within science as technological advances in physics,
chemistry and engineering attracted funding and
support. These advances also revolutionised biology,
permitting breakthroughs in physiology, biochemistry
and the molecular sciences, but in this brave new
world the diversity and distribution of organisms lost
their attraction as fields of study. It was only towards
the end of the twentieth century when environmental
issues became such a major theme in politics and with
the public that there was renewed interest in the study
of biodiversity, and a need to bring new techniques
and approaches to ‘old fields’. By then, many of the
essential skills underpinning the study of biodiversity
were already in decline. (Biodiversity itself is a word
of recent origin — first coming to the fore with the
publication of Wilson [1988].)
The taxonomic side of Linnaeus’ achievements
was outstanding. While his sexual system did not long
survive as a basis for plant classification, Linneaus had
grasped the potential for classification to be predictive
and ‘natural’, even if his particular approach had its
limitations. He had recognized the importance of the
hierarchical approach and provided a nomenclatural
system that was functional and, importantly, had
practical application to the large number of new
species that were being discovered outside Europe. It
was the first classification system that was accessible
to the non-specialist, with the work encapsulated in
handbooks that were ‘small enough to be carried into
the field’ (Koerner, 1999, p40).
In the year 2007, we celebrated the tercentenary of
Linnaeus’ birth. It also happens to be the anniversary
of Georges-Louis Leclerc, Comte de Buffon, the
author of the Histoire Naturelle. Conniff (2007), in his
article aptly titled Happy Birthday, Linnaeus, argues
that Buffon should be regarded as at least the equal
of Linnaeus, and in particular suggests that Buffon
had a superior understanding of habitat, anticipating
the development of ecology as a science. Linnaeus
and Buffon were mutually fierce critics, and Buffon
was undoubtedly also a major figure in the history of
science: he had better geological insight than Linnaeus,
and was closer to having an evolutionary perspective.
However, to suggest that Linnaeus’ natural history
LINNAEUS: KING OF NATURAL HISTORY
was not ecologically focused is far from accurate. A
major part of Linnaeus’ teaching was based on field
excursions: he had a very comprehensive knowledge
of the local flora of southern Sweden, and while
his sexual system was not appropriate for higher
taxonomic ranks, Linneaus’ species concepts have
largely stood the test of time.
We would argue that Linnaeus’ taxonomic work
was firmly underpinned by a deep understanding
of natural history and that natural history in turn
provided the basis for ecology (Mayr, 1997; Blunt,
1971). The ecological insights of Linnaeus are
clearly seen in his botanical ‘text book’, Philosophia
Botanica (Linnaeus, 1751). Koerner (1999) notes that
‘...he described many of the mechanisms of species
interdependence, as Charles Darwin noted on reading
his Oeconomia naturae of 1749’ (p15).
LINNAEUS AND NATURAL HISTORY
‘Natural history’, the advancement of which is
the prime objective of the various Linnean Societies
around the world, has a very long history. Early hunter-
gatherer societies could not have survived unless
members possessed what we might consider to be
an innate understanding of natural history, including
the ability to recognise different sorts of food and
to distinguish between the edible, the toxic and the
dangerous. Cave paintings provide, in a tangible
form, evidence for knowledge of natural history.
In classical times, plants and animals were seen as
sources of medicines or as an element in a broader
natural philosophy, and the Greeks and Romans left a
documentary record which at least in part survives to
this day and that would have been known to Linnaeus.
Knowledge of natural history would have been current
amongst the broader population, the majority of
which lived in rural environments and were intimately
dependent on the natural world for survival, but in
‘academic’ circles natural history was increasingly
associated with medicine. For hundreds of years
herbalists recycled the writings of classical authors,
without making original observations and with the
claims becoming more fanciful on each retelling. The
Renaissance then brought a new curiosity about the
world and more organized scientific inquiry, although
the importance of the links to medicine continued,
and old myths still retained currency. Linnaeus
himself was Professor of Medicine and Botany at
Uppsala University, and in some institutions close
links between the two disciplines survived until the
twentieth century. The recent growth of interest in
alternative medicine suggests a need for revitalizing
246
the links to scientific botany, and ethnobotany has
been given a new impetus as a field of study by the
regime for rewarding traditional owners of knowledge
and resources, which was established by the United
Nations Convention on Biological Diversity 1992.
From the late 16 century onwards there was
a considerable interest in collecting and studying
‘curiosities’ of all kinds, and some of the collections
of natural history objects that were assembled were
large. Some of the more academic natural historians
associated with these collections were distinguished
scientists whose work has stood the test of time. An
example is John Ray, who was the first to draw a
distinction between Monocots and Dicots and who is
commemorated in the still existing Ray Society and in
the name of the herbarium at the University of Sydney.
Some of the impetus for collecting was stimulated by
the increasing numbers of exotic specimens being
sent back to Europe from wider exploration. Linnaeus
was very much part of this natural history tradition
and although he did not travel beyond Europe, he
actively encouraged his students to do so, and he was
familiar with non-European plants both in the form
of herbarium specimens and in gardens. The non-
European species he described famously included
bananas, which would then have been regarded as
very exotic.
Linnaeus’ own exploration was closer to home
and included his early expedition to Lapland.
Although there are suggestions that his account of his
travels is somewhat exaggerated! (Koerner, 1999), it
established his reputation as an explorer and natural
historian. Lapland in the eighteenth century was at
the edge of the world and for many Europeans would
have been regarded with as much trepidation as Africa.
Even today it remains one of the few wilderness areas
in Europe (Ratcliffe 2006).
Once he was established as a senior academic
in Uppsala, field teaching became an essential and
popular part of Linneaus’ teaching. His excursions
attracted large numbers of students and were organized
with almost military precision (Blunt 1971). Most
attention was paid to the flora, although any matter
of natural history interest was open for study and
comment. Many of the localities around Uppsala
that were visited on excursions still support the same
species today, so that it requires no great stretch of the
imagination to visit sites today and see what Linneaus’
students would have seen, and to experience the same
excitement of first encountering a wet meadow full of
snakeshead fritillaries (Fritillaria meleagris) or a dry
calcareous esker with a spring abundance of Pasque
flowers (Anemone pulsatilla).
Proc. Linn. Soc. N.S.W., 130, 2009
P. ADAM AND E. MAY
LINNAEUS AND ECOLOGY
William Stearn (in Appendix I of Blunt, 1971)
recognised that Linnaeus has been variously declared
‘a pioneer ecologist, a pioneer plant-geographer, a
pioneer dendrochronologist, a pioneer evolutionist...’
but considered that the ‘most influential and useful
of his contributions to biology undoubtedly is his
successful introduction of consistent binomial specific
nomenclature’.
Itis true that Linneaus’ contribution to ecology and
plant geography is rarely acknowledged within these
disciplines, and the recognized founding fathers were
all much more recent. Nevertheless his Philosophia
Botanica contains many ecological insights, which
were in the published literature and were dormant
seeds for many decades. Given the very large number
of students who attended Linneaus’ classes, and the
wide circulation of his publications, the ecological
perspective he developed must have been assimilated
into the perceived wisdom of the day, and when
ecology and plant geography developed as separate
disciplines, Linneaus’ ideas would have been part of
the assumed background.
Today the major concerns of ecology include
the identification and evaluation of biodiversity. Of
the three generally accepted levels of biodiversity,
Linneaus was ignorant of genes, but he clearly
recognized the need to document species, and
recognized that species occupied habitats. In fact, he
devotes part of the Philosophia to discussing the main
habitats (communities) in Sweden. He also indicated
what notes should be made on field excursions.
Unfortunately, the details on many herbarium
labels in current collections fail to provide any
ecological information. Linnaeus’ advocacy of the
systematic recording of habitat data was part of his
approach to cataloguing information and these features
are easily accommodated in modern databases. If
Linnaeus were alive today, he would undoubtedly
be active in the development of bioinformatics and
the creation and manipulation of databases. The
omissions of the past cannot be corrected but today’s
collectors should be encouraged to record much more
than is often the case. Regrettably, ecologists are
often amongst the worst offenders when it comes to a
lack of detail associated with voucher specimens.
Linneaus was well aware of the variability
displayed by some species and devoted Chapter IX
of the Philosophia to a discussion of ‘Varieties’.
He urged against giving taxonomic recognition to
environmentally determined phenotypic variation,
as he recognized that a variety of diseases and insect
attack could cause abnormalities in plants (Philosophia
Proc. Linn. Soc. N.S.W., 130, 2009
section 312), displaying evidence of very careful
observation. He also pointed out that variation could
be correlated with soil conditions and microclimate
and advocated an experimental approach (Philosophia
section 316: “Cultivation is the mother of very many
varieties and is the best means of testing varieties’),
foreshadowing by a century and a half experimental
taxonomy (genecology), which enjoyed its heyday in
the second half of the 20" century.
Chapter XI of Philosophia (entitled ‘Sketches’)
contains much ecological material. Section 334 ‘The
native locations of plants relate to region, climate,
soil and ground’ contains a very succinct introduction
to ecology and biogeography (as well as some rather
strange views about geology). The discussion about
the relationship between latitude and flora gives hints
of the ideas subsequently developed in greater detail by
Alexander von Humboldt. The relationships between
soil types and the plants they support also introduce
topics that formed a major part of ecological research
in the twentieth century. Section 335 provides an
overview of phenology and indicates that Linnaeus
was well aware that factors such as temperature and
day length were involved in controlling flowering,
although it was to be many years before physiological
understanding of the mechanisms involved was
achieved. Even on botanical excursions students
recorded the plant species eaten by particular animal
species ‘while watching the botanical specimens
disappear at the moment they realized that they
needed to identify them’ (Koerner 1999 p49).
Chapter V of Philosophia (Sex) includes
observations on annual seed production of individual
plants, probably the first scientific exploration of
plant demography. The essential feature of the
Philosophia is the importance of observation, and it
is remarkable how much was achieved using lenses
and microscopes that today would be regarded as
woefully inadequate.
While Linnaeus was a creationist, the recognition
of variation suggests that he was not as rigidly so as he
is usually portrayed - he certainly recognized that the
appearance of species could change. A synthesis of
Buffon’s and Linnaeus’ ideas could have accelerated
the development of evolutionary theories, well ahead
of the publications of Darwin and Wallace in the mid
19" century.
The Philosophia is also strongly focused on
the utilisation of plants, not just as medicines but
for a whole range of purposes. The 365" (and final)
article states: ‘The economic use of plants is of great
utility to the human race.’ (Linnaeus, 1751). One of
the major justifications for biodiversity conservation
is the maintenance of the ecosystem services that
LINNAEUS: KING OF NATURAL HISTORY
biodiversity supports. This is a concept that would
clearly have found favour with Linneaus, and the
sorts of observations he advocated are needed to
document ecosystem processes. He recorded details
of the trophic interactions between organisms and had
an appreciation of the recycling of materials, noting
people used churchyard soil for growing cabbages,
hence ‘human heads ... turn into cabbage heads’
(Koerner, 1999, p83).
Robert MacArthur (1972) famously wrote that
‘to do science is to search for repeated patterns’
and stressed the importance of natural history as the
starting point for ecological research. MacArthur
pointed out that not every natural historian was
a scientist (in terms of approach and method, not
necessarily profession) and not all ecologists were
natural historians, but we would agree with him that
most of ecology has its roots in natural history: even
theoretical mathematical ecology starts with ideas that
are ultimately based on field observation. Underwood
(2007) has recently observed ‘one of the great joys
of experimental ecology is that natural history is
SO important in the development of explanatory
models’.
COLLECTING BIODIVERSITY: PRESERVING
BIODIVERSITY?
In the eighteenth, nineteenth and early twentieth
century much natural history involved collections;
and many large collections of, for example, insects,
bird eggs, or plants were made both by, or for, major
institutions and individual collectors. Many people
who subsequently became famous scientists in other
fields (for example Macfarlane Burnet - Sexton
1999) were avid collectors in their youth. Charles
Darwin himself was an avid beetle collector in his
college days (preferring ‘beetling’ to mathematics
- Desmond and Moore 1991). (Another suggestion
for Linnaeus’ mis-representation of his travels in
Lappland was mis-calculation (Selander, 1947 in
Koerner, 1999). Perhaps he shared Darwin’s aversion
to mathematics?). The making of collections taught
the need for careful observation, systematic recording
of data, and provided in-depth understanding of
particular groups of organisms.
Collectors and recorders were not just the clergy
and the landed gentry (or their spinster siblings); there
was, at least in the United Kingdom, a very strong
working class element of miners and factory workers,
who, in their very limited free time, spent many
hours completing arduous hikes and making major
finds of often taxonomically-challenging organisms.
248
This tradition of the extremely skilled amateur was
never as strong in New South Wales as it was in the
United Kingdom (for reasons that perhaps require the
attention of a social historian). Certainly, there have
been some very gifted amateurs, but their interests
tended to be restricted to groups such as birds or
flowering plants; there was not the same interest or
expertise shown in, for example, cryptogamic plants
as was the case in the United Kingdom. Given the
dearth of professionals in Australia, this means that
there are major components of biodiversity about
which we remain still basically ignorant.
Today collection is frowned upon, and in many
cases (such as the collecting of bird eggs) it is properly
illegal. The shift away from collection partly reflects
lack of opportunity given an increasingly urbanized
population, rejection of a “stamp collecting’ approach
to science and greater concern for conservation.
Certainly, it would not be possible to condone egg-
collecting or capturing and killing native vertebrates
outside specially-approved scientific licences, but
nevertheless it is probable that both school and
university students are missing out on what previously
had been important educational experiences.
The old collections remain of continuing value,
providing comparative material for taxonomic
studies, as well as evidence of changing distributions
or of environmental change. For example, the ability
to measure long-term trends in thickness of the
shells of raptor eggs was extremely important in
drawing attention to the effects of the new post-war
organic agricultural chemicals (Ratcliffe 1967, 1970;
Olsen and Olsen 1979). Sadly, curation of historical
collections in some of our greatest museums 1s being
eroded as funding for the care of collections, in the
management schemes of modern-day directors,
is coming a distant second behind promotion of
exhibitions for entertainment’s sake without the
underlying scholarship being obvious.
Even in the absence of collections, skilled
amateurs in the United Kingdom have been able
to systematically record distributions of large
numbers of taxa at the national scale. These spatially
explicit data are of enormous value for monitoring
environmental change. Data of this sort would be very
difficult to collect (and mordinately expensive) if we
had to rely on professionals, yet they will be crucial
to our monitoring of biodiversity. In Australia the
omithologists have pioneered systematic recording
of species distribution at the continental scale,
harnessing the skills and enthusiasm of amateurs and
professionals. The differences in distribution and
abundance between the two editions of the Bird Atlas
(Blakers et al. 1984; Barrett et al. 2003) provides
Proc. Linn. Soc. N.S.W., 130, 2009
P. ADAM AND E. MAY
compelling evidence for the impacts of environmental
change, and the recording program will continue into
the future. Other areas of natural history have not
been so well served.
Teaching in the field is still a component of many
University courses but is under great pressure because
of cost, occupational health and safety issues (which
can create bureaucratic nightmares), the large number
of students with part-time jobs who find it difficult to
attend courses at weekends or during vacations, and
because of the decline in the number of academic staff
with knowledge of many groups of organisms. The
long-term future of field teaching is very uncertain and
this will have major consequences for our ability to
produce graduates capable of addressing biodiversity
issues.
Peter Marren (2002, 2005) has written on
a number of occasions, pointing out the loss of
expertise in UK institutions and the decline in the
numbers of skilled amateurs. At the same time
there is an increase in the membership of NGO
conservation societies, indicating wide support for
natural history, but the deep engagement with some
particular field within natural history is less common.
There may be many and varied reasons for this, but
Marren suggests one may be that, with the modern
pressures on many people, and the absence of time,
natural history had become a spectator rather than
participant sport. The quality and expertise shown in
recent TV natural history programs 1s such that rather
than encouraging participation they suggest that we
already know everything. This is an idea that needs
further exploration.
The achievements of Linneaus and _ his
students were remarkable and the detail of their
observations made with minimal technological aids,
was particularly remarkable. Few students today
would have the capacity or patience to make similar
investigations. Experimental science needs to be
underpinned by substantial bodies of observation in
order that appropriate hypotheses can be generated
and tested. Linnaeus’ legacy of observation has been
built upon for the last 250 years, but the capacity to
continue to do so is being lost.
Contrary to the impression left by Conniff, we
would argue that Linnaeus also laid the foundation for
other major areas of ecology, including comparative
biogeography (long before van Humbolt), plant
demography, and comparative anatomy. His legacy
was far more encompassing than taxonomic, even
though the systematic recording of species remains
absolutely fundamental to modern ecological
concerns.
Proc. Linn. Soc. N.S.W., 130, 2009
REFERENCES
Barrett, G.W., Silcocks, A.F., Barry, S., Cunningham, R.D.
and Poulter R. (2003). ‘The new atlas of Australian
birds’. (Birds Australia [Royal Australasian
Ornothologists Union]: Hawthorn East, Victoria)
Blakers M., Davies S.J.J.F. and Reilly, P.N. (1984).
“The atlas of Australian birds’. (Royal Australasian
Ornithologists Union and Melbourne University
Press: Carlton, Victoria)
Blunt, W. (1971). ‘The Compleat Naturalist. A Life of
Linnaeus’. (Collins: London).
Conniff, R. (2007). Happy Birthday, Linnaeus. Natural
History 115: 42-47
Desmond, A. and Moore, J. (1991). ‘Darwin’. (Michael
Joseph: London).
Koerner, L. (1999). “Linnaeus: Nature and Nation’.
(Harvard University Press: Cambridge,
Massachusetts).
Linnaeus, C. (1751). ‘Philosophia Botanica’. (Translated
by Stephen Freer.) (Oxford University Press:
Oxford).
MacArthur, R.H. (1972). “Geographical ecology: patterns
in the distribution of species’. Harper and Row: New
York.
Mayr, E. (1997). ‘This is Biology. The Science of the
Living World’. (Belknap-Harvard University Press:
Cambridge, Massachusetts).
Marren, P. (2002). “Nature Conservation - A Review of
the Conservation of Wildlife in Britain 1950-2001’.
(Harper Collins: London).
Marren , P. (2005). “The New Naturalists- half a century of
British natural history. 2nd Edition’. (Harper Collins:
London).
Olsen, P. and Olsen, J. (1979). Eggshell thinning in the
peregrine, Falco peregrinus (Aves: Falconidae), in
Australia. Australian Wildlife Research 6, 217-226.
Ratcliffe, D.A. (1967). Decrease in eggshell weight in
certain birds of prey. Nature (Lond) 215, 208-10.
Ratcliffe, D.A. (1970). Change attributable to pesticides
in egg breakage frequency and eggshell thickness in
some British birds. Journal of Applied Ecology 7,
67-115.
Ratcliffe, D.A. (2006). ‘Lapland: a natural history’. (Yale
University Press: New Haven, Connecticut).
Selander, S. (1947). Linné 1 Lule Lappmark. Svenska
Linnéséllskapets Arsskrift. XXX, 9-20
Sexton, C. (1999). ‘Burnet, a life’. (Oxford University
Press: Melbourne).
Underwood, A.J. (2007). Logic, design, analyses,
interpretations: essentials of ecological
experimentation. British Ecological Society Bulletin
38, 24-27.
Wilson, E.O. (ed) (1988). ‘Biodiversity’ (National
Academy Press: Washington DC).
249
LINNAEUS: KING OF NATURAL HISTORY
(Endnotes)
' He variously reported traveling up to three times
250
further than he really did and spoke of long
periods spent with the native Sami, when in fact it
was a few weeks. The fact that he was being paid
by the mile for his journey by the Science Society
of Uppsala is cited by way of partial explanation
(Koerner 1999).
Proc. Linn. Soc. N.S.W., 130, 2009
Erratum
Wright, A.J. (2008). Emsian (Early Devonian) tetracorals (Cnidaria) from Grattai Creek, New South Wales.
Proceedings of the Linnean Society of New South Wales 128, 83-96.
Figure 4 on page 92 of the above paper was published without scale bars. It is reproduced here:
Ca ee Ma te a og
> ‘
~*
’
“)
oe.
Wo gs oe
Cpeyene" -.<
Sa a ee
kat
* ‘
, MMF
transverse view ; b
b)
Figure 4a-b. Trapezophyllum grattaiensis sp. nov., holotype. a, MMF 34186b
=5 mm.
34186a, longitudinal view. Bar scales
if wanotesy
viele nae sipert'n sa ihe cubes! Ae, ihe rey Kieee <P: | ie ove
ahi piso 8 OT joist reseed ect OY steidiarten pr aavon Lo a 4a008)~A
iv. (eh ile fe Wie hated dy mie AER cna sont " ie arya, Lasin bias at
CF Caen a een Pee vay ‘af rp tian Li Hiyn Ng i i
(Kerry 8 al he atiborgess mT lewd bis btw ti ai No m bit ah 9 ye !
sibstian re
EMT aft waty serovars) ORT Wh ug sian wee. ae spcalgamanti routes wean eb
Ashi ili we wooly tecibatigaal sit
INSTRUCTIONS FOR AUTHORS
(this is an abbreviated form — the full instructions can be obtained from our web site or from the Secretary)
1. The Proceedings of the Linnean Society of New South Wales publishes original research papers dealing with
any topic of natural science, particularly biological and earth sciences.
2. Manuscripts should be submitted to the Editor (M.L. Augee, PO Box 82, Kingsford NSW 2032). All
manuscripts are sent to at least two referees and in the first instance three hard copies, including all figures and
tables, must be supplied. Text must be set at one and a half or double spacing.
3. References are cited in the text by the authors’ last name and year of publication (Smith 1987, Smith and
Jones 2000). For three of more authors the citation is (Smith et al. 1988). Notice that commas are not used
between the authors’ names and the year, ‘and’ is spelled out (not &), and et al. is not in italics.
The format for the reference list is:
Journal articles:
Smith, B.S. (1987). A tale of extinction. Journal of Paleontological Fiction 23, 35-78.
Smith, B.S., Wesson, R.I. and Luger, W.K. (1988). Levels of oxygen in the blood of dead Ringtail
Possums. Australian Journal of Sleep 230, 23-53.
Chapters or papers within an edited work:
Ralph, P.H. (2001). The use of ethanol in field studies. In ‘Field techniques’ (Eds. K. Thurstle and P.J.
Green) pp. 34-41. (Northwood Press, Sydney).
Books:
Young, V.H. (1998). ‘The story of the wombat’. (Wallaby Press, Brisbane).
4. An abstract of no more than 200 words is required. Sections in the body of the paper usually include:
INTRODUCTION, MATERIALS AND METHODS, RESULTS, DISCUSSION, ACKNOWLEDGEMENTS
and REFERENCES. Some topics, especially taxonomic, may require variation.
5. Subheadings within the above sections should be in the form:
Bold heading set against left margin
This is the form for the first level headings and the first line of text underneath is indented
Underlined heading set against left margin
This is the next level, and again the first line of text underneath is indented.
Further subheadings should be avoided.
Italics are not to be used for headings but are reserved for genus and species names.
6. Up to 10 KEYWORDS are required. These are often used in computer search engines, so the more specific
the terms the better. ‘Australian’ for example is useless. Please put in alphabetical order.
7. Paragraphs are to be set off by a tab indentation without skipping a line. Do not auto-format the first line (1.e.
by using the “first line” command in WORD). All auto-formatting can be fatal when transferring a manuscript
into the publisher platform.
8. Details of setting up the manuscript:
Use 12 point Times New Roman font.
Do not justify
Margins should be: 3 cm top, 2.5 cm bottom, 3 cm left and 2.5 cm right. This is the area available for text;
headers and footers are outside these margins.
9. 3. The final version, incorporating referees’ and editor’s comments, must be supplied on floppy disc or CD in
WORD for PC format (Mac discs will not be accepted). The text file must contain absolutely no autoformatting
or track changes.
INSTRUCTIONS FOR AUTHORS
FIGURES:
Figures can be line drawings, photographs or computer-generated EXCEL or WORD files. No figures
will be accepted larger than 15.5 X 23 cm. Width of lines and sizes of letters in figures must be large enough
to allow reduction to half page size. If a scale is required, it must be presented as a bar within the figure and its
length given in the legend. It is the editor’s prerogative to reduce or enlarge figures as necessary and statements
such as “natural size” or “4X” in the legend are unacceptable.
Photographs must be supplied as black and white prints or as .TIF fi les scanned at 600 dpi. Jpeg is not
acceptable and figures of any kind set in WORD are unacceptable. Line drawings must be supplied as original
copies or as . TIF -files scanned at 1200 dpi. Other figures must be in hardcopy.
While there is no objection to full page size figures, it is journal policy to have the legend on the same
page whenever possible and figures should not be so large as to exclude the legend. Figure legends should be
placed together on a separate page at the end of the manuscript.
TABLES
Because tables may need to be re-sized, it is essential that table legends are not set within the table
but are supplied separately with the figure legends. Legends need to be the same font and size as the rest of the
manuscript.
While the text of the legend is expected to be in 12 point type, it may be necessary to use a smaller font
size for large tables. It is journal policy to accept tables that run over more than one page only in exceptional
circumstances.
Do not use vertical lines in tables unless absolutely necessary to demark data columns. Keep horizontal
lines to a minimum and never put a border around tables.
WORD or EXCEL tables are acceptable, but WORD 1s preferred.
Tables and/or figures must be separate from the text file. Never embed figures or tables in the text.
10. Details of punctuation, scientific nomenclature, etc. are to be found in the complete instructions available
from the website or from the Secretary.
11. It is helpful if authors suggest a running head of less than 40 characters.
254 Proc. Linn. Soc. N.S.W., 130, 2009
PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W.
VOLUME 130
INSTITUTION LIBRARIES
uN
SMITHSONIAN
il
wit
3 9088 01491 1994
Issued 18 March 2009
CONTENTS
PROCEEDINGS OF THE LINNEAN SOCIETY OF N.S.W.
VOLUME 130
Issued 18 March 2009
CONTENTS
21
139
Harris, J.M. and Goldingay, R.L.
Museum holdings of the broad-headed snake Hoplocephalus bungaroides (Squamata: Elapidae).
Mackness, B.S.
Reconstructing Palorchestes (Marsupialia: Palorchestidae) - from Giant kangaroo to marsupial ‘tapir’.
Choo, B.
A basal actinopterygian fish from the Middle Devonian Bunga Beds of New South Wales, Australia.
Myerscough, P.J.
Fire and habitat interactions in regeneration, persistence and maturation of obligate-seeding and resprouting plant
species in coastal heath.
Rickards, R.B., Wright, A.J. and Thomas, G.
Late Llandovery (Early Silurian) dendroid graptolites from the Cotton Formation near Forbes, New South Wales.
Chalson, J.M. and Martin, H.A.
A Holocene history of the vegetation of the Blue Mountains, New South Wales.
Chalson, J.M. and Martin, H.A.
Modern pollen deposition under vegetation of the Blue Mountains, New South Wales.
Strusz, D.L.
Silurian rhynchonellide brachiopods from Yass, New South Wales.
Wood, A.E.
Cortinarius Fr. subgenus Cortinarius in Australia.
Percival, |.G.
Late Ordovician Strophomenide and Pentameride Brachiopods from central New South Wales.
Percival, |.G.
Rare fossils (Conulata; Rostroconchia; Nautiloidea) from the Late Ordovician of central New South Wales.
Pickett, J., Och, D. and Leitch, E.
Devonian marine invertebrate fossils from the Port Macquarie block, New South Wales.
Stewardson, C.L., Prvan, T., Meyer, M.A. and Ritchie, R.J.
Age determination and growth in the male South African fur seal Arctocephalus pusillus pusillus
(Pinnipedia: Otariidae) using external body measurements.
Adam, P. and May, E.
Linnaeus: king of natural history.
Erratum.
Instructions for authors.
Printed by Ligare Pty Ltd,
138 Bonds Rd. Riverwood 2210
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11